VOL. 129, PART 1

31 MAY, 2005

Transactions of the

Royal Society of South

Australia Incorporated

INCORPORATING THE

Records of the

South Australian Museum

Contents

Watts, C. H. S. & Leys, R. Review of the epigean species of Australian Limbodessus

Guignot (Insecta: Coleoptera: Dytiscidae)

Martin, H. A. & Specht, R. L. Sclerophyll (heathy) Loins e's in the Mount Lofty

Ranges, South Australia

White, J. M. & White, T. C. R. Macro-invertebrates captured in artificial substrates in the

restored Watervalley Wetlands in South Australia — - ~— —

Horr, G., Pring, A. & Zbik, M. The Kimba meteorite: An (H4) chondrite from South

Australia

Anstis, M. & Tyler, M. J. ciel bi casey of Litoria microbelos (C ea (Anura:

Hylidae) —

Clark, P., Holz, P. & Spratt, D. M. Hepatozoon tachyglossi sp. nov.

(Haemogregarinidae), a protozoan parasite from the blood of a

short-beaked echidna, Zachyglossus aculeatus -— — - -— —-

Kemper, C. M. Records of humpback whales Megaptera novaeangliae in South

Australia — - = — -

Smales, L. R. A redescription of Odilia emanuelae (Nematoda:

Trichostrongylina: Heligmonellidae) from Australian rodents with a

key and comments on the genus Odilia

Tanner, J. E. Three decades of habitat change in Gulf St Vincent, South Australia

Souter, N. J. Flood regime change in the Hattah Lakes Victoria resulting from

regulation of the River Murray =

Brief Communication:

O’Callaghan, M., Reddin, J. & Lehmann, D. Helminth and protozoan parasites of feral

cats from Kangaroo Island =

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

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

Transactions of the Royal Society of S. Aust. (2005), 129(1), 1-13.

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT

(INSECTA: COLEOPTERA: DYTISCIDAE).

by C. H. S. WatTs* & R. Leyst

Summary

watts, C. H. S. & Leys, R. (2005). Review of the epigean species of Australian Limbodessus Guignot (Insecta:

Coleoptera: Dytiscidae). Trans. R. Soc. S. Aust. 129(1), 1-13, 31 May, 2005.

DNA sequencing of the CO1 and 16s-tRNA-NDI fragments of the mitochondrial genome was used to support

the morphological results. Nine species are recognised; Limbodessus amabilis (Clark), L. capeensis sp. nov.,

L. compactus (Clark), L. gemellus (Clark), L. inornatus (Sharp), L. occidentalis (Watts & Humphreys),

L. praelargus (Lea), L. shuckardii (Clark), and L. rivulus (Larson). Limbodessus dispar (Sharp) is synonymised

with L.' shuckardii (Clark). A key to the species is provided.

Key Worbs: Coleoptera, Dytiscidae, Limbodessus, Taxonomy, DNA.

Introduction

Among the commonest diving beetles in southern

Australia are members of the genus Limbodessus

Guignot of the tribe Bidessini, which are often

abundant in still to slightly moving, shallow water.

The genus level classification of these beetles has

recently been revised by Balke and Ribera (2004),

who, based on morphological and DNA sequence

data, synonymised the genera Boongurrus Larson,

Tjirtudessus Watts and Humphreys, the Australian

members of Liodessus Guignot and Limbodessus

Guignot, placing them all in Limbodessus. Cooper et

al. (2002) had previously studied DNA sequence

data of species in these genera and tentatively come

to similar conclusions but made no taxonomic

decisions regarding them.

The species level taxonomy of the species

formally placed in Tjirtudessus, Boongurrus and

Limbodessus has recently been dealt with by Watts

and Humphreys (2003), Larson (1994) and Balke

and Sato (1995) respectively. No similar revision has

been done for the six Australian species previously

placed by Watts (1978) in Liodessus. Increased

collecting since 1978 has highlighted the great

similarity between these species and_ the

consequential difficulties in their identification.

The need for a soundly based taxonomy of these

species has become more urgent with the realisation

that they are congeneric with the numerous

subterranean species, previously placed in

Tjirtudessus, being discovered in Western and

Central Australia (Watts & Humphreys 1999, 2003,

“ Department of Entomology, South Australian Museum, North

Terrace, Adelaide, South Australia 5000.

t Evolutionary Biology Unit (and Centre of Evolutionary Biology

and Biodiversity) South Australian Museum, North Terrace,

Adelaide, South Australia 5000

2004). To understand the evolution of these

stygobitic species a better understanding of the

taxonomy of the epigean members of the genus is

needed.

This paper mainly deals with the species level

taxonomy of those Australian species of Bidessini

previously classified as Liodessus (Watts 1978,

2002). Utilising the extensive collection in the South

Australian Museum, it is based on adult morphology,

in particular the male genitalia, supported by DNA

sequence analysis of the CO1 and 16S-tRNA-NDI

fragments of the mitochondrial genome of key

specimens. For details of the DNA procedures used

see Cooper ef al. (2002). For completion, brief notes

are given on the three other epigean Limbodessus

species that were previously placed in Limbodessus

and Boongurrus (Watts 2002; Watts and Humphreys

2004).

Unless otherwise noted all specimens were

collected by C. H. S. Watts.

Abbreviations

BMNH The Natural History Museum, London.

SAMA South Australian Museum, Adelaide.

TABLE |. Observed DNA sequence divergence among

Limbodessus species previously placed in Liodessus.

Comparison % Sequence divergence

Within the Limbodessus spp. 2.9-10.8

given below.

Between sister species:

L. inornatus (3) - L. gemellus (3) 2.9-3.7

L. amabilis (1) - L. praelargus (1) 353.

Within species:

L. gemellus (3) 0.1-0.3

L. shuckardii (1)/L. dispar (3) 0.0-0.7

(n) = number of sequenced specimens per species.

Results

The combined morphological and DNA results

recognise six species, one of which, L. capeensis, is

new.

Limbodessus dispar (Sharp) from the Southwest

is morphologically indistinguishable from the

eastern L. shuckardii (Clark). DNA sequence data

confirm a close relationship with specimens from

Western Australia and Victoria differing at only

0.0% — 0.7% sequence divergence (Table 1). We

consider them to be the same species.

Specimens morphologically identified as

L. gemellus (Clark) (see keys and under species

descriptions) from Tasmania (19 km W. Maydeena),

Adelaide (Onkaparinga gorge) and the Flinders

Ranges (Moro Gorge) are closely similar

biochemically (Table 1). These differ by 2.9 — 3.7%

sequence divergence from specimens identified as L.

inornatus (Sharp) from the Southwest. These are

morphologically close but differ in the distal shape of

the penis. We consider that the sequence and

morphological differences are sufficient to consider

them separate species. Limbodessus amabilis (Clark)

and L. praelargus (Lea) are morphologically

indistinguishable other than by their differently

shaped penises. The sequence data confirm their

specific separation (Table 1).

Systematics

Diagnosis

The specimens discussed here share the following

morphological features.

Epigean Australian Bidessini. Elongate-oval,

without sutural striae, with well-developed elytral

and pronotal plicae, with long metacoxal lines,

paramere two segmented, with well-developed

finger-like apical lobe on distal segment of paramere

(Figs 11-18). (Subterranean species of Limbodessus

are more varied in shape and may lack the elytral and

pronotal plicae and metacoxal lines (Watts and

Humphreys 2003).)

Key to epigean species of Limbodessus

Including Allodessus bistrigatus (Clark) which is

closely related (Balke and Ribera 2004) and is often

confused with species of Limbodessus.

1. Extreme front of elytral epipleuron with raised

transverse carina delineating a basal pit...............

Pen ee ners pier pare Limbodessus compactus (Clark)

— Elytral epipleuron without basal pit... 2

2. Head with fine line between back edges of eyes

(GCeOrviCal Strid)... ccscissssccnssccavsessscsccessenesssesvaseets 4

— Head without cervical stria or, if present, only

weakly and partially (Pilbara, Yilgarn and

Atherton Tableland )........ccccccccsssceccseesesseeeesnees 3

C.H. 8. WATTS & R. LEYS

Eyes of normal size, hind edges reaching well

beyond anteriolateral corners of pronotum.

Western Australia (Fig. 26)........... L. occidentalis

poten heWeatedsahog sadertanenbte badcetante (Watts & Humphreys)

Eyes reduced to about 60% normal size, hind

edges not, or only just, reaching anteriolateral

corners of pronotum. Atherton Tableland (Fig.

QEY-cteschacoeddeartpenvenleclonsersieedosegt L. rivulus (Larson)

Paramere with apical segment simple, finger-like

(Fig. 11). Dorsal surface reddish/yellow to

greyish, elytra with diffuse darker markings;

ventral surface in female with black metathorax

and metacoxae and yellow abdomen, males black

except prosternum and head; segments of female

antenna with a basal groove; elytra densely

but relatively weakly punctate, coxal plates

moderately punctate ......0 cece eee eeeeeeeeneereeetees

or thptes Desh sthoiehs cleesae Allodessus bistrigatus (Clark)

Paramere with apical segment strongly lobed,

often hook-like (Figs 13-20). Segments of female

antennae without basal groove; ventral surface

usually more uniformly coloured; punctures

Watia ble: assesssizeescrtceyssdicrzeceracns\nteescanssnapsot eae 5

Metacoxal plates with numerous large punctures;

(Northeast Queensland, Fig. 22)...

Saaedgedabiens tebe outed ateaecneade L. capeensis sp. nov.

Metacoxal plates virtually impunctate ............... 6

Pronotal plicae straight; never with dorsal surface

mat; antenna relatively thin (Fig. 1); pro and

mesotarsi in males moderately expanded (Fig. 5):

paramere with apical lobe without setae (Fig. 13).

SOUHSAS Aree seid. oh resi carstieemntaeresaqeegpanents any 7

Pronotal plicae slanted or curved inwards; may

have mat dorsal surface; antenna, particularly in

females, stout (Figs 3, 4); pro and mesotarsi

strongly expanded, particularly in the males (Figs

7, 8); paramere with apical lobe with long setae

(Fig. 20). Southeast and Southwest... 8

Penis with apical quarter relatively broad,

noticeably narrowing only close to tip (Fig. 13)..

stale deste boL Qs bob uted MeTah sgh badh sled iaeiieny L. amabilis (Clark)

Penis with apical quarter narrowing to blunt point

(Fig, 1B) nits snceiatteeneansbiebens L. praelargus (Lea)

Penis suddenly narrowing to narrow apical

quarter (Fig. 20); length 2.5 — 3.0 mm; pronotum

with disc usually diffusely darker; elytron with

colour pattern tending linear; pronotal plicae tend

to curve inwards in apical half; antenna,

particularly in female, stout (Fig. 4)... eee

spaamiehau nea eanibig ate necks aathad enna be L. shuckardii (Clark)

Penis evenly narrowing to tip in apical quarter

(Figs 15, 16); length 2.2 — 2.8 mm; pronotum

never with disc diffusely darker; elytron with

colour pattern blotchy; pronotal plicae sinuate;

antennae not as stout (Fig. 3)... eee eee 9

Penis with apical portion long and thin, parallel

sided (Fig. 15); protibiae in female moderately

& eb ¢

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT

-—— —

g a

Figs 1-4. Male and female antennae of: 1, Limbodessus amabilis; 2, L. capeensis; 3, L. inornatus; 4, L. shuckardii.

Figs 5-8. Male and female protarsi of: 5, Limbodessus amabilis, 6, L. capeensis, 7, L. inornatus; 8, L. shuckardii.

4 C. H.S, WATTS & R. LEYS

broad (Fig. 10); Southeast (Fig. 24)...

HO Gestadeth ers yeetlofttconmt tbh eels L. gemellus (Clark)

Penis with apical portion smoothly narrowing to

tip (Fig. 16); protibiae in female very broad (Fig.

9); Southwest (Fig. 25)........ L. inornatus (Sharp)

Descriptions.

(In alphabetical order.)

Limbodessus amabilis (Clark)

Hydroporus amabilis Clark, 1862, p. 420.

Liodessus amabilis (Clark). Guignot, 1939, p. 54.

Limbodessus amabilis (Clark).

Balke and Ribera, 2004.

Types

Lectotvpe (designated by Watts 1978). Upper

specimen of two mounted on separate cards on same

pin, “amabilis Clark” (yellow label), no data,

BMNH.

Paralectotypes 1, pinned under lectotype,

“amabilis Clark” (yellow label); 3 females, | male

mounted on same card, “67.56/ amabilis Clark”; 5,

“S. Australia” “Bakewell 59,24”. All BMNH.

Description (number of dissected males examined,

27). Figs 1, 5, 13, 21.

Habitus. Length 2.6 — 3.3 mm; moderately

convex; not constricted at junction of

pronotum/elytra; narrowly oval.

Head. Dark reddish, often lighter towards front.

Narrower than elytra. Smooth, shiny, punctures small

but deep, rather sparse, stronger and denser towards

the rear; cervical stria well marked. Antenna with

segment | cylindrical, segment 2 as long as segment

1, barrel-shaped, segment 3 as long as segment 2,

narrower, narrowing towards base, segment 4 shorter

than segment 3, segments 5-10 subequal, segment | |

about twice length of segment 10, narrower (Fig. 1).

Maxillary palpus elongate, segment 4 as long as

segments 1-3 combined.

Pronotum. Reddish-yellow, diffusely darker

towards rear in some. A little narrower than elytra;

anteriolateral angles projecting strongly forward;

base not constricted, posteriolateral angles square,

surface shiny, punctures relatively sparse, uneven in

size, small on disc, larger behind, particularly

inwards from plicae; basal plicae strong, excavated

somewhat on inside, straight, reaching about half

way to front margin of pronotum.

Elytra. Dark reddish, most specimens with

indistinct light/dark pattern, occasionally distinct.

Elongate, widest at middle; shiny, moderately

densely and somewhat unevenly covered with

moderate punctures; plicae well impressed, straight,

about as long as pronotal plicae. Elytron with well

developed inner ridge near apex (ligula). Epipleuron

lacking basal carina, relatively broad in anterior

quarter, then progressively narrowing to near apex.

Ventral surface. Meso and metathorax, metacoxae

and abdomen reddish-brown to black, appendages,

pronotum and head lighter. Prosternal process

narrow between coxae, reaching mesothorax, apical

half relatively broad with parallel raised ridges on

each side, not arched in lateral view. Metathorax with

wings short, broadly rounded in midline behind, with

scattered very small punctures. Metacoxal plates

large, shiny; punctures very small, scattered;

metacoxal lines raised, distinct, moderately widely

spaced, reaching to metasternum, weakly diverging;

closely adpressed to ventrite 1. Ventrites | and 2

fused, with a few very large deep punctures, sutural

line distinct, ventrites 3 to 5 mobile, with a few small

punctures, somewhat denser and larger on ventrite 5.

Legs. Protibia triangular, outer edge straight or

weakly bow-shaped, widest at apex where it is about

four times its basal width; protarsus weakly expanded,

segment | about 2x as long as broad, segment 2 a little

wider than segment 1, about half as long, segment 3

about as long as segment 1 and same width, deeply

bifid, segment 4 very small, hidden within lobes of

segment 3, segment 5 narrow, cylindrical, a bit longer

than segment 3, segments | to 3 with dense covering

of adhesive setae (Fig. 5); claws short and simple.

Mesotrochanter elongate, subrectangular, with a row

of relatively long thin setae on inner edge; mesofemur

with 2 to 3 moderately strong setae near base on hind

margin, stronger than those on mesotrochanter,

mesotarsus slightly longer than protarsus.

Metatrochanter tip rounded; metafemur elongate,

lacking spines; metatibia narrow, moderately curved,

widening towards apex; metatarsus relatively stout,

segment | longest, segment 5 longer than segment 4,

segments | and 2 in combination about as long as

others; claws weak.

Male.

Little external difference between the sexes except

that antennae in the female are a little stouter (Fig. 1).

Median lobe of aedeagus moderately broad, apex

rounded (Fig. 13). Paramere quite broad, apical

segment with apical lobe overlapping apex of rest of

segment which is broadly rounded (Fig. 13).

Remarks

A common species in South Australia, Victoria and

Tasmania, often occurring together with L. gemellus,

L. shuckardii or L. praelargus. Indistinguishable

from L. praelargus other than by the broad tip to the

penis, it is separable from the other two species by its

relatively unexpanded pro and mesotarsi and straight

rather than curved or sinuate pronotal plicae. Never

with a mat female form such as occurs in L.

shuckardii and L. gemellus.

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT 5

The lectotype is a female and hence, on present

knowledge, cannot be separated from L. praelargus.

Among the paralectotypes is an immature male. The

teneral penis appears more like that of L. praelargus

than L. amabilis (M. Balke pers. com.) suggesting that

some of the paralectotypes are in fact L. praelargus.

We have however retained the usage of Watts 1978 and

associated the name L. amabilis with this species.

Specimens examined (localities of males with

genitalia extracted).

New South Wales. Berry 1/68; 14 km W Delegate,

4/11/97. Vietoria. Buangor, 9/11/97; 1 km S Drik

Drik 11/10/97; Mt Emu Creek, 1/59; 3 km S

Fisherman’s Rest, 6/11/97. South Australia.

Adelaide, 11/60; Hindmarsh Island, 8/62; Chain of

Ponds, 12/62; 3 km W Yunti, 5/10/95; Mt Compass,

8/61; 10 km E Mt Compass, 10/9/97; 1 km S$

Nangwarry, 5/10/00; Williamstown, 10/61.

Tasmania. 25 km E Bridport, 23/1/00; 5 km E

Bridport, 23/1/00; Cradle Valley Cradle Mountain-

Lake St Clair NP, 19/1/00; Ellendale, 1/12/00, 2 km

SW Ellendale, 2/12/00; 2 km W Fingal, 23/1/00;

Swansea Jan; Haartz Mt, 15 km W_ Geeveston,

3/12/00; Harcus River, 14 km SW Montagu,

22/1/00; Hobart, Jan; 12 km N Hobart, 2/12/00; 8 km

N Kingston, 3/12/00.

Limbodessus capeensis sp. nov

Types

Holotype

Male. “Mcllwraith Rng Weather Stn N. Qld

23/7/82. C.Watts.” SAMA, SAMA Data Base # 25-

002891.

Paratypes

4, as for holotype, SAMA, SAMA Data base # 25-

009337; 4, “Captain Billy Ck N.Q. 10/83 C. Watts.”,

SAMA, SAMA Data base # 25- 009338.

Description (number examined 9). Figs 2, 6, 14, 22.

Habitus. Length 1.9 — 2.1 mm; moderately convex;

very weakly constricted at junction of pronotum

/elytra; narrowly oval.

Head. Reddish-yellow. Narrower than elytra.

Smooth, shiny, punctures small but deep, rather

sparse; cervical stria well marked. Antenna with

segment | cylindrical, segment 2 as long as segment

1, barrel shaped, segment 3 as long as segment 2,

narrower, narrowing towards base, segments 4-10

becoming progressively slightly broader, segment 1 1

about twice length of segment 10, narrower (Fig. 2).

Maxillary palpus elongate, segment 4 as long as

segments 1-3 combined.

Pronotum. Reddish-yellow. A little narrower than

elytra; anteriolateral angles projecting strongly

forward; base very weakly constricted, poster-

iolateral angles square, surface shiny, punctures

relatively sparse, uneven in size, small on disc, larger

behind, particularly inwards from plicae; basal plicae

strong, excavated somewhat on inside, sinuate,

nearly reaching front margin of pronotum.

Elytra. Rather uniformly dark reddish with tips

lighter, some specimens with indistinct light/dark

pattern. Elongate, widest behind middle; shiny, quite

densely and evenly covered with strong punctures;

plicae well impressed, straight, sloping inwards a

little, about as long as pronotal plicae. Elytron with

well developed inner ridge near apex (ligula).

Epipleuron lacking basal carina, relatively broad in

anterior quarter, then progressively narrowing to near

apex.

Ventral surface, Meso and metathorax, metacoxae

and abdomen reddish-brown, appendages, pronotum

and head lighter. Prosternal process narrow between

coxae, reaching mesothorax, apical half relatively

broad with parallel raised ridges on each side, not

arched in lateral view. Metathorax with wings short,

broadly truncated in midline behind, with large deep

punctures. Metacoxal plates large, shiny; punctures

moderately dense, large, deep; metacoxal lines

raised, distinct, moderately widely spaced, reaching

to metasternum, weakly diverging; closely adpressed

to ventrite 1. Ventrites | and 2 fused, with moderate

number of very large deep punctures, sutural line

distinct, ventrites 3 to 5 mobile, sparsely covered

with small punctures, somewhat denser and larger on

ventrite 5.

Legs. Protibia triangular, outer edge weakly bow-

shaped, widest at apex where it is about four times its

basal width; protarsus very weakly expanded,

segment | about 2x as long as broad, segment 2 a

little wider than segment 1 and a little shorter,

segment 3 as long as segment | and a little broader,

deeply bifid, segment 4 very small, hidden within

lobes of segment 3, segment 5 narrow, cylindrical,

about same lengtli as segment 3, segments | to 3 with

dense covering of adhesive setae (Fig. 6); claws short

and simple. Mesotrochanter elongate, sub-

rectangular, with a few relatively long thin setae on

inner edge; mesofemur with 2 to 3 moderately strong

setae near base on hind edge, much stronger than

those on mesotrochanter, mesotarsus slightly longer

than protarsus. Metatrochanter tip rounded;

metafemur elongate, lacking spines; metatibia

narrow, moderately curved, widening towards apex;

metatarsus relatively stout, segment | longest,

segment 5 longer than segment 4, segments | and 2

in combination about as long as others; claws weak.

Male

Little external difference between the sexes.

Median lobe of aedeagus moderately broad in

6 C.H.S. WATTS & R. LEYS

middle, narrowing in apical quarter to blunt point

(Fig. 14). Paramere elongate, apical segment with

stout apical lobe well separated from rest of segment

(Fig. 14).

Remarks

Very similar to some Australian Leiodytes in size,

colour and strong punctation. Differs from Leiodytes

in the lack of a slightly raised margin to the front of

the head and in the lobed parameres. The small size,

unexpanded pro and mesotarsi and very strong

ventral punctures readily separate this species from

all other Australian Liodessus. The apical lobe of the

paramere is less well developed than in other

Australian species. Known only from Cape York.

Etvmology

From its locality — Cape York.

Limbodessus compactus (Clark)

Hydroporus compactus Clark, 1852, p. 421.

Bidessus compactus (Clark). Sharp, 1882, p. 362.

Limbodessus compactus (Clark).

Guignot, 1939, p. 53.

= Bidessus neoguineensis Regimbart.

Balke & Sato, 1995, p. 188.

= Uvarus tokarensis Sato.

Balke & Sato, 1995, p. 188.

Limbodessus compactus is the type species of the

genus. It has recently been comprehensively dealt

with by Balke and Sato (1995) and because of this is

not redescribed here.

Diagnosis

1.6 — 2.5 mm long, boat-shaped, stout antennae,

shiny, uniformly reddish, relatively convex, with

cervical stria, with basal carina on elytral epipleuron,

male genitalia as in Fig.12.

Remarks

The species is widely distributed in Japan, SE Asia,

New Guinea and Australia (Balke & Sato 1995) (Fig.

23). It is readily separated from other Limbodessus

species by its uniform reddish colour, boat-like shape

and the presence of transverse carina at the base of

the elytral epipleura. Within Australia it is

widespread (see below), favouring the littoral zone of

still water. In pools in sandy riverbeds in inland

Australia the species has been collected interstitially

for a short distance beyond the edge of freestanding

water. Limbodessus occidentalis and L. rivulus occur

in similar places but penetrate further interstitially

than L. compactus.

Specimens examined

New South Wales. 1, Armidale, 1/61; 1, ditto,

21/3/63; 1, Barrington, 17/8/97; 25, 2 km N

Batemans Bay, 2/11/97; 2, Berry, 1/63; 5, Collector,

2/61; 2, 2 km N Collector, 26/11/98; 10, 8 km N

Failford, 18/8/97; 3, Maclean, 1/61; 3, Nyngan,

16/3/63; Smith’s Lake, 5/70. Northern Territory. 1,

1 km SE Batchelor, 12/4/66, N McFarland; 2,

Cahills crossing, Kakadu National Park, 9/6/73,

Upton & Feehan; 25, Cannon Hill Kakadu National

Park, 10/10/98; 1, 10 mi E Daly River, 26/6/72, BK

Head; 3, Darwin, 13/5/63; 1, Gubara, Kakadu

National Park, 12/10/98; 1, Howard Springs,

23/3/98; 2, Jabiru, 12/10/98; 3, Mt Borradaile,

8/10/98: 1, 4 mi W Timber Creek, 14/4/66, N

McFarland. Queensland. 1, Alligator River, 20 km

S Townsville, 25/3/96; 1, 8 km N Bluewater,

3/11/95; 1, Brisbane, 1/61; 17, Bushland Beech, 20

km N Townsville, 26/2/98, AJ. Watts; 3, ditto,

15/3/98: 1, ditto 16/1/98; 4, ditto, 6/2/98; 4,

Caloundra, 27/3/63; 5, ditto, 7/3/63; 1, Cairns,

16/4/63; 1, Greenvale, 27/3/96; 2, 70 km SW

Greenvale, 28/3/98, AJ Watts; 1, ditto, 11/3/96; 2,

Mackay, 4/4/63; 2, Malanda, 13/4/63;

1, 5 km NW Mt Molloy, 5/2/97; 4, Nardello’s

Lagoon, 29/3/96; 2, Stanthorpe, 1/61; 2, 25 km §

Townsville, 25/3/96; 4, 37 km S Townsville,

6/11/95; 1, Townsville, 31/10/95. South Australia.

1, Naracoorte, 9/3/71; 1, 25 km NE Mt Gambier,

26/3/82, FA. Forrest. Tasmania. 4, 6 km N Pioneer,

13/1/00. Victoria. 2, Dartmoor, 12/61; 7, Fern Tree

Gully, 12/61; 6, Healesville, 12/68; 1, 12 km SW

Orbost, 30/11/98; 1, 12 km SW Orbost, 5/11/973; 1,

Stratford 7/11/97. Western Australia. 2, Cane River

HS (old), 22/5/01; 1, Cane River, 22/5/01; 1, 12 km

S Newman, 27/5/01; 4,1 km N Red Hill stn.,

22/5/01; 3, 30 km N Red Hill stn., 22/5/01; 8,

Wittenoom Gorge, 26/5/01.

Limbodessus gemellus (Clark)

Hydroporus gemellus Clark, 1862, p. 421.

Bidessus gemellus (Clark). Sharp, 1882, p. 362.

Liodessus gemellus Clark). Watts, 1978, p. 49.

= Bidessus mundus Sharp, 1882, p. 362.

Watts, 1978, p. 49.

Limbodessus gemellus (Clark).

Balke and Ribera 2004.

Types

Bidessus gemellus Clark.

Lectotype “gemellus Clark Australia” (yellow

label), BMNH.

Paralectotypes. 2, “gemellus Clark” (yellow label),

BMNH.

Bidessus mundus Sharp.

Lectotype. ‘Type 88/Australia / n.sp. australia /

Bidessus mundus/ (male symbol)”, BMNH.

Paralectotypes. 1, “Type 88/australia”; 1,

Australia”; both BMNH.

“88 /

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT Zt

VE.

Figs 9-10. Male and female protibia of: 9, Limbodessus

inornatus; 10, L. shuckardii.

Description (number examined, 125.) Figs 15, 24.

As for L. amabilis except as follows.

Habitus. Length 2.5 — 3.0 mm; weakly constricted

at junction of pronotum and elytra.

Head. Reddish/yellow, darker towards rear.

Pronotum. Reddish-yellow, diffusely darker along

front boarder; basal plicae strong, excavated

somewhat on inside, curving inwards slightly,

straightening towards front.

Elvira. Elongate, widest at middle; moderately

densely and evenly covered with moderate

punctures; plica well impressed, straight, slanting

inwards slightly, about as long as pronotal plicae.

Legs. Protibia bow-shaped, widest near apex where

it is about four times its basal width, protarsus

moderately expanded (cf. Fig. 7).

Male

Dorsal surface shiny; antennae, tarsi and protibiae

relatively stout (cf. Figs 3, 7, 9). Median lobe of

aedeagus broad at base, narrowing to long, thin,

apical portion (Fig. 15). Paramere broad, apical

segment weakly triangular, apical lobe large,

overlapping rest of apical segment at base, with fine

setae (Fig. 15).

Female

Dorsal surface duller, often with weak to moderate

reticulation. Antennae a little stouter. Protibia wider

and more triangular (cf. Fig. 9); pro and mesotarsi

slightly less expanded (cf. Fig. 7).

Remarks

Similar in size to L. amabilis and L. praelargus but

more uniformly coloured, with reticulate females,

sinuate pronotal plicae, broad pro and mesotarsi and,

in the female, broad pro and mesotibiae and stout

antennae. Separable from the slightly smaller L.

shuckardii by the lack of a darker disc on the

pronotum, the pronotal plicae more sinuate, and the

absence of vague linear markings on the elytra.

Limbodessus inornatus from the Southwest is very

similar morphologically but differs in having a

longer, parallel-sided distal portion to the penis

(Fig. 16) and female L. inornatus have somewhat

broader pro and mesotibiae (Fig. 9). With a more

inland distribution than L. amabilis and L. praelargus

but not to the same extent as L. shuckardii.

Specimens examined

New South Wales. 3, 2 km N Batemans Bay,

18/4/97; 1, Berry, 1/68; 1, 3 km N Bulli, 27/11/98; 2,

8 km N Failford, 18/8/97; 1, Gosford 1/61; 1,2 km S

Nowra, 27/1/00; 3, 2 km S Nowra, 27/11/98:

1, Quaama, 18/1/97; 2, Ulladulla, 2/1/973;

1, Windsor, Lea; 2, Waterfall, 1/6/82. Queensland. 1,

Queensland, Blackburn coll. South Australia. 1,

Adelaide, Griffith coll: 1, Chain of Ponds, 10/11/96; 1,

Dalhousie Spr. at light, 6/10/87, J.A.Forrest; 1,

Flinders Range, May 59; 1, 10 km N Forreston,

17/9/96; 1, Kuipo 35° 14’ § 145° 138’ E, 5/10/95; 4,

Moro Gorge, Flinders Ranges R. Leys; 13, Mt

Gambier, 12/16; 3, Myponga, 9/12/96; 3, 13 km W

Meadows 36° 11’ S 138° 96’ E, 28/9/96; 2, 10 km E

Mt Compass, 10/9/97; 1, Mt Lofty, Lea; 2, Myponga,

5/1/88; 3, 1 km S Nangwarry, 9/10/97; 3, Port Lincoln,

Blackburn; 3, ditto, 1/82; 1, Wood Point River Murray,

4/11/80, P. Waller. Tasmania. 1, 5 km E Bridport,

23/1/00; 1, Cradle Mt N P, Jan; 1, 17 km SW Derwent

Bridge, 29/11/00; 3, 18 km N Derwent Bridge,

24/1/00; 1, Ellendale, 1/12/00; 1, 2 km SW Ellendale,

1/12/00: 1, 5 km SE Gormanston, 29/11/00; 1,

Hatfield River, 15 km SW, 28/11/00; 3, Hobart, 8/61;

2, Hobart, Griffith; 3, 12 km N Hobart, 2/12/00; 1,

King Isl, collector unknown; 1, 8 km W Kingston,

3/12/00; 1, 9 km Maydena, 1/12/00; 11, Narcissus Bay

Lake St Clair, 30/11/00; 1, 6 km N Pioneer 23/1/00; 3,

8 C.H.S. WATTS & R. LEYS

2 km W Port Latta, 27/11/00; 1, St Helens, Jan; 1,

Swansea, Jan; 1, 16 km N Waratah, 28/11/00; 1, 9 km

N Queenstown on B28 Road, 28/12/00. Victoria. 1,

Buangor, 9/11/97; 4, Dartmoor, 11/10/97; 2, 17 km

SW Derwent Bridge, 29/11/00; 2, Healesville, 12/68;

3, 8 km W Kingston, 3/12/002; 1, Omeo, 6/11/97; 3,

30 km W Portland, 10/10/97; 1, Sardine Ck 30 km, N

Orbost, 16/1/97; 4, Stratford, 7/11/97.

Limbodessus inornatus (Sharp)

Bidessus inornatus Sharp, 1882, p. 360.

Liodessus inornatus (Sharp). Watts, 1978, p. 48.

= Bidessus biformis Sharp, 1882, p. 362.

Watts, 1978, p. 48.

Limbodessus inornatus (Sharp).

Balke and Ribera, 2004.

Types

Bidessus inornatus Sharp.

Holotype

Male. “Type 81/West Australia/ Bidessus inornatus

n. sp’’, BMNH.

Bidessus biformis Sharp

Lectotype. “W. Australia/ type 89 d Bidessus

biformis n.sp. K.G.Sound”, BMNH.

Paralectotypes. 3, same data as lectotype, BMNH.

3, “W. Australia, Swan River” BMNH. 6, “W.

Australia”, BMNH.

Description (number examined, 351) Figs 3, 7, 9,16,

25:

As for L. gemellus except as follows.

Male

Median lobe of aedeagus relatively broad,

narrowing in apical quarter to blunt tip (Fig. 16).

Paramere broad, apical segment rounded at apex,

apical lobe long, overlapping rest of segment, with

fine setae and some peg-like structures on front edge

(Fig. 16).

Female

Pro and mesotibiae expanded (Fig. 9).

Remarks

Restricted to the Southwest where it is sympatric

with L. shuckardii. Can be separated from this

species by its larger size, uniform colour on the

pronotum, and the pronotal plicae sinuate rather than

curved. Female L. inornatus have slightly broader

pro and mesotibiae (Fig. 9). The front of the apical

lobe of the paramere has several peg-like structures

not found in other Australian Limbodessus (Fig. 16).

Indistinguishable from the eastern L. gemellus apart

from a stouter distal portion to the penis and the peg-

like structures on the parameres.

Specimens examined

Western Australia. 3, Armadale, 3/62, D.

Edwards; 3, Bickley Swamp, Rottnest Island, 10/58,

D. Edwards; 1, Blackwood River, Nannup, 20/10/96;

1, Bonanup, 17/10/96; 13, Bridgetown, 11/9/31,

Darlington; 3, 50 km W along Broke inlet Rd Nr

Walpole, 18/9/00; 4, 3 km N Bullsbrook, 16/10/96;

1, Bushy Swamp 15 km WNW _ Woodanilling,

21/9/00; 16, Byenup Lagoon NR, 21/9/00; 6, Corio

Spring Rottnest Island, 10/58, D. Edwards; 4, 5 km

W Cowaramup, 22/10/00; 1, 2 km SW Dandalup,

23/9/00; 2, 20 km SE Donnybrook, 18/10/96; 3, 30

km S Dwellingup, 17/10/96; 1, 8 km S Dwellingup,

17/10/96; 1, Ellen Brook NR, 14/9/00; 8, 19 km S

Fremantle, 24/10/96; 1, Geraldton 10/31, Darlington;

Hay River, collector unknown; 13, Ironstone Gully

Falls 13 km SW Donnybrook, 22/10/96; 15, 10 km E

Kalamunda, 16/10/96; 7, Kodjinup NR, 21/9/00 : 7,

Lake Pleasant View, 17/9/00; 1, Lake Parkeyerring,

15/9/00; 1, Lake Poorginup, 20/9/00; 17, Maidavale,

27/4/90; 11, Margaret River, 10/31, Darlington; 7, 4

km N Mumballup, 23/9/00; 1, Nalyerin Lake,

22/9/00; 13, 4 km S New Norcia, 15/10/96; 15 km

NW Pemberton, 17/5/87; 1, 20 km W Pemberton,

20/10/96; 7, Pemberton, 10/31, Darlington; 2, 15 km

NE Pemberton, 8/10/96; 69, 6 km S_ Pinjarra,

23/9/00; 48, 30 km N Perth, 14/10/96; 1, Rottnest

Island, 10/31, Darlington; 3, 12 km W Serpentine,

24/10/96; Stirling Range, 10/72; 2, Swan River, Lea;

9, Swan River, 10/57, D. Edwards; 11, 20 km W

Strachan, 21/9/00; 1, 1 km S Wagin, 21/9/00; 1,

Wilgarup River, 6/58, D. Edwards; 1, 16 km N

Woodanilling, 15/9/00; 4, Yallingup, 22/10/96.

Limbodessus occidentalis (Watts and Humphreys)

Boongurrus occidentalis

Watts and Humphreys, 2004.

Limbodessus occidentalis (Watts & Humphreys)

This species has only recently been described

(Watts and Humphreys 2004) and hence will not be

redescribed here.

Diagnosis

1.9 — 2.3 mm long, relatively flat, elongate-oval,

light reddish, elytra darker, without cervical stria,

without basal carina on elytral epipleuron, eyes of

normal size, male genitalia as in Fig. 17.

Remarks

Limbodessus occidentalis differs from all other

epigean Limbodessus except L. rivulus, by the

absence of a cervical stria between the hind edges of

the eye. Like L. rivulus it is small (1.9 — 2.3 mm

long), flattened with a rugose surface but unlike L.

rivulus the eyes are not reduced in size. It occurs at

the headwaters of sandy/gravely streams, or at the

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT 9

GOP G

Figs 11-18. Dorsal view of penis, lateral view of penis and paramere of: 11, Allodessus bistrigatus; 12, Limbodessus

compactus; 13, L. amabilis; 14, L. capeensis; 15, L. gemellus; 16, L. inornatus; 17, L. occidentalis; 18, L. praelargus.

10 C. H. S. WATTS & R, LEYS

upstream edges of pools in drying riverbeds, in the

Pilbara and Yilgarn regions of Western Australia

(Fig. 26), often interstitially several meters away

from the water’s edge. It has also occasionally been

taken together with a true stygobitic fauna in shallow

calcrete aquifers (Watts and Humphreys 2004).

Specimens examined (Holotype & Paratypes)

Western Australia. 9, 10 km NW Eerala Stn,

23/5/01; 1, Killara Station, 6/6/02,W. F. Humphreys

& R. Leys); 3, Moorarie Station, 8/6/02, W. F.

Humphreys & R. Leys; 1, Wagga Wagga Station,

4/6/02, W. F. Humphreys & R. Leys; 12, Wittenoom

Gorge, Town Pool, 26/5/01, C. H. S. Watts &

G. A. Watts.

Limbodessus praelargus (Lea)

Bidessus praelargus Lea, 1898, p. 523.

Liodessus praelargus (Lea). Watts, 1978, p. 51.

Limbodessus praelargus (Lea).

Balke and Ribera, 2004.

Types

Holotype

Male, “praelargus Lea TYPE 6d Forest Reefs”;

dissected and remounted this study, SAMA. SAMA

data base # 25-001525.

Paratypes

5 females, same locality; dissected and remounted

this study, SAMA.SAMA data base # 25-009153.

Description (number of dissected males examined,

13) Figs 18, 27.

As for L amabilis except:

Male

Medial lobe of aedeagus moderately broad,

smoothly narrowing to a blunt point (Fig. 18).

Paramere broad, apical lobe stout, overlapping

apical portion of apical segment which is relatively

narrow and has the apex pointed rather than rounded

(Fig. 18).

Remarks

Indistinguishable from L. amabilis except for the

pointed penis and narrower paramere. The range of

the two species is broadly similar. Watts (1978)

correctly (but fortuitously) associated the name

praelargus with the species with the pointed penis.

See also under L. amabilis.

Specimens examined (localities of males with

genitalia extracted.)

Australian Capital Territory. 30 mi S Canberra,

1/61. New South Wales. 14 km W Delagate, 4/11/97.

Victoria. | 1 km E Bruthen, 6/11/97; 4 km S Glenista,

24/9/98; 10 km NE Mirranatwa, 12/10/97; 5 km NW

Portland, 10/10/97. South Australia. Mt Gambier,

12/61; 1 km S Nangwarry, 9/10/97. Tasmania. S end

of Lake St Clair, 30/11/00; Little Pine Lake, 8 km W

Miena, 23/10/00; 2 km W Port Latta, 27/11/00.

Limbodessus rivulus (Larson)

Boongurrus rivulus Larson, 1994.

Limbodessus rivulus (Larson).

Balke and Ribera, 2004.

This species has recently been treated in detail

(Larson 1994) and hence will not be redescribed here.

Diagnoses

1.8 — 2.2 mm long, rather rectangular, relatively

flat, dull reddish with darker areas, strongly

reticulate, lacking (or virtually lacking) cervical

stria, lacking basal carina on elytral epipleuron, eyes

much smaller than normal for epigean Limbodessus,

male genitalia as in Fig. 19.

Remarks

Together with L. occidentalis the only epigean

Limbodessus lacking a cervical stria between the rear

of the eyes although Larson (1994) reported its

partial presence in a small number of specimens.

Separated from L. occidentalis by its obviously

reduced eyes and characters of the male genitalia

(Watts and Humphreys 2004). The species occurs at

(

Eee

Figs 19-20. Dorsal view of penis, lateral view of penis and paramere of: 19, Limbodessus rivulus; 20 L. shuckardii.

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT

he A

OP Se TF

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27 ‘eg!

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ate “yeu y é r: i \y

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Figs 21-29. Distribution maps of specimens of Limbodessus in the South Australian Museum Entomological collection. 21,

28, L. rivulus; 29, L. shuckardii.

L. amabilis; 22, L. capeensis: 23, L. compactus; 24, L. gemellus; 25, L. inornatus: 26, L. occidentalis; 27, L. praelargus;

12 C.H.S. WATTS & R. LEYS

the headwaters of small, gravely, spring-fed streams

in areas of open eucalypt woodland on the Atherton

Tablelands. Here it occurs in gravel at the edge of the

water or interstitially for at least a metre from the

waters edge (pers. observation). Very recently

(August, 2004), I have collected a single specimen of

this, or a closely related species, from similar habitat

on Mt Tamborine in southeast Queensland.

Specimens examined

Queensland. |, | km E Watsonville, 31/3/96; 11,

1.5 km E Watsonville, 3/8/03.

Limbodessus shuckardii (Clark)

Hydroporus shuckardii Clark, 1862, p. 420.

Bidessus shuckhardii (Clark). Sharp, 1882, p. 361.

Liodessus shuckhardi (Clark). Watts, 1978, p. 47.

Limbodessus shuckardii (Clark).

Balke and Ribera, 2004.

Bidessus dispar Sharp, 1882, p. 363.

Liodessus dispar (Sharp). Watts, 1978, p. 48.

= Bidessus elegans Lea, 1898, p. 523. Watts 1978,

p. 48.

Limbodessus dispar (Sharp). Balke and Ribera,

2004.

=Limbodessus shuckardii (Clark). Syn. nov.

Types

Hydroporus shuckardii Clark

Lectotype. *67.56 /shuckardii Clark australia’

(yellow label), BMNH.

Paralectotypes. 1, ‘shuckardii (yellow label); 2, ‘S.

Australia. Bakewell, 59.24; 10, ‘australia’; all in

BMNH.

Bidessus dispar Sharp.

Lectotype. ‘Type 90 2 a / Swan River / W.

Australia/ W. Australia / Sharp Coll 1905-313 /

Bidessus dispar Sharp 2 a Type’ (in Balfour-

Browne’s writing).

Paralectotypes. 2, same locality; 1, ‘W. Australia’;

both in BMNH.

Bidessus elegans Lea.

Holotype

‘elegans Lea TYPE Beverley’; left-hand specimen

on card marked with TY, SAMA. SAMA data base #

25-001582.

Paratype

Same data as Holotype; right-hand specimen on

card, SAMA. SAMA data base # 25-009340.

Description (number examined, 244.) Figs 4, 8,10,

20, 29.

As for L. amabilis except:

Habitus. Length 1.9 — 2.6 mm; very weakly

constricted at junction of pronotum and elytra.

Head, Light reddish-yellow with darker patches

towards rear. Antenna (male) stout, segment |

cylindrical, segment 2 as long as segment 1, barrel-

shaped, segment 3 as long as segment 2, narrower,

narrowing towards base, segment 4 shorter than

segment 3, segments 5-10 subequal, slightly wine-

glass shaped, segment 11 about twice length of

segment 10, narrower (Fig. 4).

Pronotum. Light reddish-yellow with diffuse

reddish-brown area on disc. Basal plicae strong,

excavated somewhat on inside, curving inwards quite

strongly, reaching about two-thirds of way to front

margin of pronotum.

Elytra. Light reddish/yellow to reddish yellow with

reddish brown pattern, 2-3 thin longitudinal lines

partially discernible in most specimens. Elongate,

widest at middle; shiny, moderately densely and

evenly covered with moderate punctures; plicae well

impressed, slanting inwards, about as long as

pronotal plicae.

Ventral surface. Light reddish-yellow, meso and

metathorax and metacoxae darker. Metathorax with

wings short, truncated or broadly rounded in midline

behind, virtually impunctate.

Legs. Protibia bow-shaped, widest before apex

where it is about 3x its basal width; protarsus

moderately expanded, segment | a little longer than

wide, segment 2 a little wider than segment 1, about

half as long, segment 3 about as long as segment |

and same width, deeply bifid, segment 4 very small,

hidden within lobes of segment 3, segment 5 narrow,

cylindrical, a bit longer than segment 3, segments |

to 3 with dense covering of adhesive setae.

Male

Shiny, antenna and tarsi as above (Figs 4, 8).

Median lobe of aedeagus moderately broad,

narrowing a little quite abruptly in apical quarter in

both dorsal and lateral views (Fig. 20), Paramere

moderately broad, apical segment rather small, apical

lobe relatively long, overlapping rest of apical

segment, with long setae (Fig. 20).

Female

Both dorsal and ventral surfaces weakly to

moderately reticulate. Antenna stouter, segments 5-

10 almost bead-like (Fig. 4). Pro and mesotarsi

broad, but less so than in male (Fig. 8).

Remarks

A relatively small species recognised by the broad

pro and mesotarsi, pronotal plicae curving inwards

and the centre of the pronotum diffusely slightly

darker than the rest. The elytra have indistinct thin

REVIEW OF THE EPIGEAN SPECIES OF AUSTRALIAN LIMBODESSUS GUIGNOT 13

linear markings reminiscent of those of

Hydroglyphus grammopterus (Zimmerman) which

can be faintly seen on most specimens. Other

Australian Limbodessus can have quite marked linear

elytral markings but these are broader and

interrupted in the central region. A more inland

distribution than other species, present as far north as

the Mount Isa — Greenvale region of north

Queensland, seemingly absent from Tasmania.

Specimens examined

New South Wales. 1, Collector, 20/1/97; 1, Forbes,

15/3/63; 2, Gilgandra, 19/11/92; 2, Ditto, 9/2/62; 2,

Grenfell, collector unknown; 9, Nyngan, 16/3/63.

Queensland. 11, Camooweal, 30/4/93; 2, Charters

Towers, 23/4/63; 1, Cloncurry, 29/4/63; 1,

Coorabulka, 7/71; 1, 70 km SW Greenvale, at light,

28/3/95 to 7/4/95, A. J. Watts; 1, ditto, 21-31/10/95; 1,

ditto, 29/1/97 to 4/2/97; 1, ditto, 3-10/10/96; 1, Lake

Buchanan, 25/9/83, B Timms. South Australia. 2,

Adelaide; 1, Alligator Gorge, 6/58; 4, Chain of Ponds,

12/62; 1, 20 km N Coober Pedy, 2/68; 2,10 km N

Forreston, 3/9/99; 1, Leigh Creek, Blackburn’s coll:

2, | km S Nangwarry, 9/10/97. Victoria. 1, Albury,

16/7/89, P. Waller; 1, 12 km W Brimpaen, 23/9/98; 5,

Buangor, 9/11/97; 1, Dartmoor, 24/9/98; 2, Dodswell

Bridge, 10/10/98, D. Churches; 12, Fyans Creek, 15

KS Stawell, 13/1/97; 2, 4 km S Glenista, 24/9/96; 6,

Grampians, 2/63; Halls Gap, 13/1/97; 11, Healesville,

12/68; 1, Lake Hattah, Light trap, 28/10/67, G. W.

Anderson; 8, 12 km N Mirranatwa, 12/10/97; 2, 10

km NE Mirrantwa, 12/10/97; 1, 5 km NW Portland,

10/10/97; 3, Nathatia, 9/6; 2, Turret Falls, 5 km NW

Halls Gap, 13/1/97; 1, Wellington River 4 km N

Licola, 30/11/98. Western Australia. 2, Armadale,

7/62, D. Edwards: |, Beverley, Lea; 8, Bridgetown,

9/11/31, Darlington; 1, 5 km N Bushy Swamp nr

Wagin, 21/9/00; 2, Byenup Lagoon NR, 21/9/00; 4, 8

km S Dwellingup, 17/10/96; 69, 6 km S§ Pinjarra,

23/10/96; 2, 8 km N Pinjarra, 23/10/96; 1, Ironstone

Gully Falls, 13 km SW Donnybrook, 22/10/96; 4, 10

km S Fremantle, 24/10/96; 1, 4 km N Mumballup,

23/10/96; 1, Nalyerin Lake, 22/9/00; 1, 5 km E Lake

Nalyerin, 22/9/00; 1, Riffle Range Swamp, Rottnest

Island, 10/59, D. Edwards; 16, 12 km W Serpentine,

24/10/96; 1, 10 km S Yallingup, 22/10/96.

Acknowledgments

We would like to thank Howard Hamer for drawing

the illustrations, Debbie Churches and Archie

McArthur for helping with the manuscript and

Micheal Balke for information on the types in the

BMNH and comments on the manuscript.

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Regimbart, from underground waters in Australia. Records

of the Foust Australian Museum 36, 135-187.

, 2004. Thirteen new Dytiscidae

(Galopiens) of the genera Boongurrus Larson,

Tjirtudessus Watts & Humphreys and Nirripirti Watts &

Humphreys, from underground waters in Australia.

Trans. R, Soc. S. Aust. 128, 99-129,

Transactions of the Royal Society of S. Aust. (2005), 129(1), 14-24.

SCLEROPHYLL (HEATHY) UNDERSTOREYS IN THE

MOUNT LOFTY RANGES, SOUTH AUSTRALIA

by H. A. Martin* & R. L. SPECHT

Summary

Martin, H. A. & SPECHT, R. L. (2005) Sclerophyll (heathy) understoreys in the Mount Lofty Ranges, South

Australia. Trans. R. Soc. S. Aust. 129(1), 14-24, 31 May, 2005.

Floristic patterns in the sclerophyll (heathy) understoreys, which are characteristic of the nutrient-poor soils of

the Mount Lofty Ranges, are surveyed: — on Pre-Cambrian schists at Inglewood (rainfall c. 800 mm per annum);

on Pre-Cambrian quartzites at Morialta (rainfall c. 800 mm per annum); on Pre-Cambrian quartzites in Waterfall

Gully (rainfall c. 1150 mm per annum); on truncated laterites of Early Tertiary in National Park Belair (rainfall

c. 800 mm per annum); and on deep Mid-Tertiary sands at Blewitt Springs (rainfall c. 800 mm per annum).

Seven Floristic Groups have been defined objectively using Goodall’s ‘positive interspecific correlation’

technique, adapted for computer-analysis. The floristic patterns appear to be determined by the annual cycle of

available soil water at each site and the slightly different nutrient levels of sandy versus clayey surface soils.

Key Woros: Floristic Groups, heath; sclerophyll, understorey, objective classification, nutrient-poor soil.

Introduction

Two distinct formations, termed Sklerophyllen-

Wald and Savannen-Wald by Diels (1906), are

characteristic of the vegetation of the Mount Lofty

Ranges (Adamson & Osborn 1924; Specht & Perry

1948: Specht er a/. 1961). The former formation,

translated as ‘dry sclerophyll forest’ (now heathy

open-forest, Specht 1970), is invariably associated

with extremely nutrient-poor soils. The latter

formation, translated as ‘savanna forest/woodland’

(now grassy open-forest/woodland, Specht 1970) is

on more fertile, but not nutrient-rich, soils (Specht &

Rundel 1990; Specht & Specht 1999). In the Mount

Lofty Ranges, the transition from one formation to

the other is determined by the geological substrate

that changes abruptly over a short distance (Sprigg

1946).

Remnants of the very infertile, lateritic soils that

were formed on the Late Cretaceous to Early Tertiary

peneplain persisted until the uplift of the Mount

Lofty Ranges and still overlie the geological

formations on the Eden-Moana Fault Block

(Stephens 1946, 1971; Specht & Perry 1948).

The original understorey of the savanna (grassy)

woodlands on the more fertile soils was dominated

by Danthonia (Adamson & Osborn 1924; Davies &

Sim 1931; Wood 1937), but is now invaded by many

introduced species (Specht & Perry 1948; Specht er

al. 1961; Specht 1972, 1975; Specht & Clifford

1991; Specht 2000, 2001, 2002). The nutrient-poor

“School of Biological, Environmental and Earth Sciences,

University of New South Wales, Sydney, New South Wales 2052.

‘ Emeritus Professor of Botany, The University of Queensland.

Current address: 107 Central Avenue, St Lucia, Queensland 4067.

soils that support sclerophyll (heathy) open-

forests/woodlands are unsuitable for the establish-

ment of invasive plants, unless the soil phosphate

levels are increased after bush-fires or from pollution

(Specht 1963; Heddle & Specht 1975: Specht &

Specht 1989, 1999; Specht 2001, 2002).

The floristics of many stands of heathy open-

forests in the Mount Lofty Ranges were collated in

the ecological survey conducted in 1946-47 (Specht

& Perry 1948) and again in ‘The Vegetation of South

Australia’ (Specht 1972). In the late 1950s, Helene

Martin examined in detail the distribution of the

component species in the gradient from heathy

woodland to heathy open-forest on Pre-Cambrian

schists (34° 48’ S, 138° 48’ E) about 2.5 km north-

east of Inglewood, north of the Torrens Gorge

(Martin 1961; Martin & Specht 1962).

In this paper, the changes in floristics of

sclerophyll (heathy) understorey vegetation are

examined in detail on the nutrient-poor soils that

have developed along the rainfall isohyet of c. 800

mm per annum:

(1) Under Eucalyptus obliqua — E. goniocalyx syn.

E. elaeophora (+ E. fasciculosa) open-forest on

Pre-Cambrian schists north-east of Inglewood

(Specht e¢ al. 1961; Martin 1961);

(2) Under E. obliqua — E. baxteri (+ E. fasciculosa)

open-forest on Pre-Cambrian quartzites at

Morialta (Specht & Perry 1948);

(3) Under E. microcarpa syn. E. odorata — E.

leucoxylon woodland on truncated laterites of the

Early Tertiary in National Park Belair (Specht &

Perry 1948);

(4) Under E. fasciculosa woodland on deep Mid-

Tertiary sands at Blewitt Springs (Specht &

Perry 1948),

SCLEROPHYLL VEGETATION IN THE MOUNT LOFTY RANGES 15

i <=

As sores

me ===

\ = eae be

Rivet

ST. VINCENT GULF

DISTRIBUTION

FORMATIONS

=| Dry sclerophy!!

= Banksia ornata

= =| dominant

~ES) Banksia marginata

- t

ns dominant

C] Savanna

oO i 3 83

a a |

Fig. |. Location of study sites. Morialta (M), Waterfall Gully (WG), National Park Belair (NPB) and Blewitt Springs (BS),

in the sclerophyll (heathy) vegetation of the Mount Lofty Ranges (after Specht & Perry 1948). The study site at Inglewood

is 12 km NNE of Morialta (Specht ef al. 1961; Martin 1961). Banksia Ornata and Banksia marginata are dominants in

the heathy understorey, not the overstorey.

16 H. A. MARTIN & R. L. SPECHT

Group 1

(726)

+ Hibbertia stricta

+ Acacia myrtifolia

Group 4 Group 5

(208) (442)

+ Astroloma conostephioides + Leptospermum myrsinoides

Group 8 Group 9 Group 6 Group 7

(82) (126) (254) (188)

+ Platylobium + Hibbertia

obtusangulum acicularis

+ Epacris impressa

Group 10 Group 11

(104) (150)

Group 14 Group 15 Group 12 Group 13

(62) (64) (141) (47)

Fig. 2. Association analysis of sclerophyll (heathy) understoreys in the Mount Lofty Ranges.

Blewitt Springs — Group 2; Inglewood — Groups 10, 11, 12, 13 and 15; Morialta — Groups 8 and 13; National Park Belair

— Groups 12 and 15; Waterfall Gully — Group 14.

SCLEROPHYLL VEGETATION IN THE MOUNT LOFTY RANGES 17

TABLE |. Major floristic composition of sclerophyll (heathy) understoreys in Floristic Groups (Fig. 2) in the Mount Lofty

Ranges. Species that enabled a dichotomous split in the ‘association analysis’ (Fig. 2) are shown in bold.

Group 2 — Blewitt Springs (42 spp.)

Tall shrubs (M): Acacia pycnantha

Low shrubs (N): Acacia myrtifolia, A. spinescens, Allocasuarina muelleriana, Banksia ornata, Calytrix tetragona,

Dillwynia sericea, Hakea ulicina, Leptospermum myrsinoides, Olearia ramulosa, Xanthorrhoea semiplana

Sub-shrubs (Ch): Astroloma conostephioides, A. humifusum, Hibbertia stricta, H. virgata

Graminoids (H): Lepidosperma carphoides

Group 4(8) — Morialta (61 spp.)

Low shrubs (N): Acacia myrtifolia, Epacris impressa, Hakea rostrata, Ixodia achillaeoides, Leptospermum myrsinoides,

Pultenaea daphnoides, Xanthorrhoea semiplana

Sub-shrubs (Ch): Acrotriche serrulata, Astroloma conostephioides, Hibbertia sericea, Leucopogon virgatus,

Tetratheca pilosa

Graminoids (H): Lepidosperma semiteres

Group 6(10) — Inglewood (39 spp.)

Low shrubs (N): Dillwynia hispida, Hakea rostrata, Leptospermum myrsinoides, Platylobium obtusangulum, Pultenaea

largiflorens, Xanthorrhoea semiplana

Sub-shrubs (Ch): Acrotriche serrulata, Astroloma humifusum, Hibbertia acicularis, H. sericea, Leucopogon virgatus,

Pimelea linifolia

Graminoids (H): Lepidosperma semiteres

Group 6(11) — Inglewood (44 spp.)

Low shrubs (N): Allocasuarina muelleriana, Hakea rostrata, Leptospermum myrsinoides, Xanthorrhoea semiplana

Sub-shrubs (Ch): Acrotriche serrulata, Astroloma humifusum, Hibbertia acicularis, H. sericea

Graminoids (H): Lepidosperma semiteres

Group 7(12) —Inglewood and National Park Belair (43 spp.)

Tall shrubs (M): Acacia pycnantha

Low shrubs (N): Pultenaea daphnoides, Xanthorrhoea semiplana

Sub-shrubs (Ch): Acrotriche serrulata, Astroloma humifusum, Hibbertia acicularis, H. sericea

Graminoids (H): Lepidosperma semiteres

Group 7(13) — Inglewood and Morialta (48 spp.)

Tall shrubs (M): Acacia pycnantha

Low shrubs (N): Olearia ramulosa

Sub-shrubs (Ch): Astroloma humifusum, Hibbertia sericea

Group 9(14) — Waterfall Gully (47 spp.)

Low shrubs (N): Acacia myrtifolia, Acrotriche fasciculiflora, Epacris impressa, Hakea rostrata, Hakea ulicina, Ixodia

achillaeoides, Leptospermum myrsinoides, Platylobium obtusangulum, Pultenaea daphnoides, Xanthorrhoea semiplana

Sub-shrubs (Ch): Acrotriche serrulata, Leucopogon virgatus, Tetratheca pilosa

Graminoids (H): Lepidosperma semiteres, Lomandra fibrata

Group 9(15) — Inglewood and National Park Belair (54 spp.)

Low shrubs (N): Acacia myrtifolia, Leptospermum myrsinoides, Platylobium obtusangulum, Pultenaea daphnoides,

Xanthorrhoea semiplana

Sub-shrubs (Ch): Astroloma humifusum, Hibberta acicularis, H. sericea

Graminoids (H): Lepidosperma semiteres

The floristics of the sclerophyll (heathy) Methods

understorey vegetation on Pre-Cambrian quartzites

at Morialta (annual rainfall c. 800 mm) and under the The location of the study sites at Morialta (34° 50’

E. obliqua — E. baxteri (+ E. cosmophylla) open- 8, 138° 40’ E), Waterfall Gully (35° 00’ S, 138° 40’

forest in Waterfall Gully (rainfall c.1150 mm per _ E), Belair National Park (35° 02' S, 138° 38’ E) and

annum) are compared. Blewitt Springs (35° 10’ S, 138° 34’ E) are shown on

18 H. A. MARTIN & R. L, SPECHT

Fig. 1 (after Specht & Perry 1948). The study site at

Inglewood (34° 48’ S, 138° 47’ E) is 12 km NNE of

Morialta.

At each site, the species composition in quadrats

(20m x Im) was recorded along. transects,

established on both north- and south-facing slopes

and spaced 20m along the top of the ridge.

Nomenclature of the species follows Jessop &

Toelken (1986).

In all, 852 quadrats were recorded: 414 on Pre-

Cambrian schists at Inglewood; 117 on Pre-

Cambrian quartzites at Morialta; 141 on Pre-

Cambrian quartzites at Waterfall Gully; 89 on

truncated laterites of the Early Tertiary in Belair

National Park; and 91 on deep Mid-Tertiary sands at

Blewett Springs. The data was collated in 1960.

Species densities (per 20m? quadrat) were plotted

for major species in the understorey at each site to

construct isonome maps (Pidgeon & Ashby 1942;

Rayson 1957; Brewer née Pidgeon 1995). The

isonome maps for the large data-set of 414 quadrats

in the sclerophyll (heathy) vegetation in the climatic

gradient north-east of Inglewood were published by

Martin (1961); the smaller data-sets at the other sites,

where rainfall is relatively uniform, are less variable

in floristic patterns.

Floristic Groups in the large data-bank of 852

quadrats (20m x Im), collected from the five

sclerophyll (heathy) sampling sites, were defined

objectively by ‘association analysis’, using Goodall’s

‘positive interspecific correlation’ technique

(Goodall 1953), adapted for computer analysis by W.

T. Williams of C.S.I.R.O. Division of Computing

Research (Clifford & Stephenson 1975). Firstly, the

chi-squared values for every pair of species in the

data-bank are calculated. Next, the sum of all the chi-

squared values for each species is totalled; the

species with the highest chi-squared sum is

considered as the ‘indicator species’ for the positive

group of the dichotomous split of the data-bank.

These positive and negative floristic groups are

considered to be distributed in different micro-

habitats in the vegetation under study (Clifford &

Specht 1979; Specht & Specht 1999),

Results

The large data-bank of 852 quadrats (20m x Im)

recorded in the sclerophyll (heathy) understoreys of

the Mount Lofty Ranges (Appendix 1), from

Inglewood, north of the Torrens Gorge, to Blewitt

Springs in the south, was analysed objectively by the

classification program ‘association analysis’ (Fig. 2

and Table 1). The objective analysis recognized eight

Floristic Groups:

Floristic Group 14, with Acacia myrtifolia,

Leptospermum myrsinoides, Pultenaea daphnoides,

and Epacris impressa common in the understorey, is

a relatively uniform community developed in high

rainfall on the quartzitic outcrop between Mount

Lofty Summit and Waterfall Gully.

Floristic Group 8, with Acacia myrtifolia,

Leptospermum myrsinoides, Astroloma cono-

stephioides, and Hibbertia sericea prominent in the

understorey, occupies most of the quartzitic outcrop

above the Waterfalls at Morialta.

Floristic Groups 10, 11, 12, 13 and 15 form a

graded series of communities from the wettest to the

driest part of the Inglewood District on podsolic soils

developed on Pre-Cambrian schists (Martin 1961).

The following species are either the most frequent

(or the most indicative) in the understorey:

Floristic Group 11 — Hakea — rostrata,

Leptospermum myrsinoides, Hibbertia acicularis, H.

sericea, Lepidosperma semiteres.

Floristic Group 12 — Platylobium obtusangulum,

Lepidosperma semiteres (+ Pultenaea daphnoides,

Lomandra fibrata)

Floristic Group 10 — Hibbertia acicularis, H.

sericea, Pimelea spathulata (+ Pultenaea

largiflorens).

Floristic Group 15 — Hibbertia acicularis,

Lepidosperma semiteres.

Floristic Group 13 — Hibbertia sericea (+ Acacia

pycnantha).

The Floristic Groups 13 and 15 in the driest part of

the sclerophyll continuum at Inglewood tend to

approach the herbaceous understorey of the grassy

savanna. The sclerophyll understorey in the driest

part of Morialta is similar to Floristic Groups 13

found at Inglewood.

The sclerophyllous understoreys on the truncated

lateritic podsol in Belair National Park belong to the

same Floristic Groups 12 and 15 found in the

Inglewood study.

Floristic Group 2, with Banksia ornata,

Xanthorrhoea semiplana, and Hibbertia stricta

widespread in the understorey, is found on the deep

sandy podsols near Blewitt Springs.

Discussion

Association analysis of the 852 quadrats (20m x

Im in dimensions) recorded in the sclerophyll

(heathy) vegetation over 40 km distance north-south

in the Mount Lofty Ranges indicated that the

Floristic Groups found on the Pre-Cambrian

quartzite of Waterfall Gully (Group 14) and at

Morialta (Group 8) were distinctly different from the

Floristic Groups found on the Mid-Tertiary sands at

Blewitt Springs (Group 2) and in the rainfall gradient

on Pre-Cambrian schists at Inglewood (Groups 10 to

13 and 15). The ecotonal Floristic Group 13 between

sclerophyll and savanna vegetation at Inglewood,

SCLEROPHYLL VEGETATION IN THE MOUNT LOFTY RANGES 19

north of the Torrens Gorge, was also found at the

driest margin of Moritalta to the south. Both Floristic

Groups 12 and 15 on Pre-Cambrian schists at

Inglewood also occurred on the truncated lateritic

podsol near Pines Oval in Belair National Park.

The shrubs, Acacia myrtifolia, Epacris impressa,

Hakea_ rostrata, Leptospermum — myrsinoides,

Pultenaea daphnoides and Xanthorrhoea semiplana

were common on the Pre-Cambrian quartzites in

Waterfall Gully (Group 14) and at Morialta (Group

8). These shrubby species were associated with sub-

shrubs such as Acrotriche serrulata, Hibbertia

sericea, Leucopogon virgatus and Tetratheca pilosa,

while Astroloma conostephioides became prominent

in the drier Morialta understorey (Group 8). The

tussock sedge Lepidosperma semiteres was common

in the ground stratum in both Groups.

Acacia myrtifolia, Allocasuarina muelleriana,

Banksia ornata, Leptospermum myrsinoides and

Xanthorrhoea semiplana, with Hibbertia stricta as a

diagnostic sub-shrub species, dominated the

sclerophyll understorey on the deep sands at Blewitt

Springs (Group 2). This understorey Floristic Group

is very similar to the treeless heathland that

dominates the deep sands in the Ninety-Mile Plain

(Specht & Rayson 1957; Rayson 1957; Specht eg al.

1958).

In the Floristic Groups on the clayey soils of the

Pre-Cambrian schists at Inglewood, the wettest

Groups (Groups 10, 11 and 15) are dominated by

shrubby species such as Hakea rostrata,

Leptospermum myrsinoides and Xanthorrhoea

semiplana, together with sub-shrubs Acrotriche

serrulata, Astroloma humifiusum, —Hibbertia

acicularis and H. sericea, with the tussock sedge

Lepidosperma semiteres in the ground stratum.

Floristic Group 15, with distinctive species such as

Acacia myrtifolia, Platylobium obtusangulum and

Pultenaea daphnoides that occur in the wetter

section of the rainfall gradient at Inglewood, is also

defined in the sclerophyll understorey on the clayey

truncated lateritic soils in Belair National Park.

The golden wattle, Acacia pycnantha occurs as

scattered small trees in the ecotone between the

sclerophyll (heathy) and the savanna (grassy)

vegetation at Inglewood, Morialta and Belair

National Park (Groups 12 and 13), where shrubby

species are rare, but sub-shrubs such as Astroloma

humifusum and Hibbertia sericea persist.

Ecological interpretation of the objective

classification, in relation to the climate and soils of

the Mount Lofty Ranges, was summarised for ‘The

Vegetation of South Australia’ (Specht 1972) and

expanded in this paper.

Conclusions

The distribution of species in the sclerophyll

(heathy) understorey of the Mount Lofty Ranges is

determined by a combination of (1) the annual cycle

of available soil water (Martin & Specht 1962;

Rayson 1957; Specht 1957a, 1957b) and (2) the

interaction of infertile sandy surface soils versus

clayey surface soils (Specht 1988; Specht & Rundel

1990; Specht & Specht 1999).

The ability of the sclerophyll (heathy) vegetation to

survive, since the Late Cretaceous, on such infertile

soils, and to be replaced by savanna (grassy)

understorey on more fertile soils, was the major

stimulus for the long-term nutrition experiments on

Dark Island heath (Specht 1963; Heddle & Specht

1975; Specht & Specht 1989, 1999),

Acknowledgements

We are indebted to Ms Paulene Riessen, then of the

Botany Department, University of Adelaide, for

assistance with field work. Dr W. T. Williams of

C.S.1.R.0. Division of Computing Research

analysed the quadrat data in 1970,

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APPENDIX TABLE |. Sclerophyll (heathy) understorey in the Mt Lofty Ranges, South Australia.

Floristic Groups: Group 2— Blewitt Springs; Group 4(8) — Morialta; Group 4(9) — Waterfall Gully, Inglewood and National

Inglewood; Group 7(12) - Inglewood and National Park Belair;

Group 7(13) - Inglewood and Morialta; Group 9(14) — Waterfall Gully; Group 9(15) - Inglewood and National Park Belair:

Life form: MM Mesophanerophyte; M Microphanerophyte; N Nanophanerophyte; Ch Chamaephyte; H Hemicryptophyte

Park Belair; Group 6(10) — Inglewood; Group 6(11)

(semi-evergreen)

Statistics: Number of quadrats (20m x Im) that were recorded containing the species, with percentage of the total number

of quadrats in parentheses below

Species Life Group 4 Group 6 Group 7 Group 2 Group 9

form 4(8) 4(9) 6(10) 611) 712) 713) 2 9(14) 915)

Asteraceae

Ixodia achillaeoides N 6l 33 20 11 23 5 - 25 8

(74) (26) (19) (9) (16) (11) (40) (13)

Olearia ramulosa N 15S 13 Il 6 5 21 65 7 6

(18) (10) (1) (4) (4) (45) (86) (11) (9)

Casuarinaceae

Allocasuarina verticillata M 8 6 - 3 2 10 - 2 4

(10) (5) (2) (1) (21) (3) (6)

Species

A. muelleriana

Cyperaceae

Life

form

Lepidosperma carphoides H

L. laterale

L. semiteres

L. viscidum

Dilleniaceae

Hibbertia acicularis

H. sericea

H. stricta

H. virgata

Epacridaceae

Acrotriche fasciculiflora

A. serrulata

Astroloma conostephioides N

A. humifusum

Epacris impressa

Leucopogon rufus

L. virgatus

Lissanthe strigosa

Fabaceae

Daviesia leptophylla

D. ulicifolia

Dillwynia sericea

D. hispida

Eutaxia microphylla

Platylobium obtusangulum N

Pultenaea daphnoides

P. involucrata

P. largiflorens

SCLEROPHYLL VEGETATION IN THE MOUNT LOFTY RANGES 21

Group 4 Group 6 Group 7 Group 2 Group 9

4(8) 4(9) 610) 611) 712) 7013) 2 914) 9(15)

21 27 35 75 19 1 43 12 15

(26) Ql) (34) (50) (14) (2) (57) (19) (23)

10 7 - 1 - 3 51 6 1

(12) (6) (1) (6) (67) (10) (2)

2 - - 1 - - 18 - -

(2) (1) (24)

44 94 101 133 103 9 - 48 46

(54) (75) (97) (89) (73) (19) (78) (72)

- / 1 - - 2 - 4 3

(6) (1) (4) (6) (5)

13 65 102 121 141 - - 17 48

(16) (52) (98) (81) = (100) (27) (75)

72 81 97 135 73 35 - 13 64

(88) (64) (93) (90) (52) (75) (21) (100)

- 1 z - - - 76 - 1

(1) (100) (2)

3 2 - - - - 45 - 2

(4) (2) (59) (3)

6 45 1 3 24 4 - 31 14

(7) (36) (1) (2) (17) (9) (50) (22)

46 72 78 100 58 6 16 41 3

(56) (57) (75) (69) (AID (13) Q1) (66) (4)

81 - 1 10 6 13 58 - -

(99) (1) (7) (4) (28) (76)

32 43 50 87 61 21 50 17 26

(39) (34) (48) (58) (43) (45) (66) (27) (41)

46 62 1 9 4 - e 62 2

(56) (49) () (6) (3) (100)

1 I 2: - 1 - 21 : 1

(1) (1) (2) (1) (28) (2)

54 43 45 45 18 5 - 26 17

(66) (34) (43) (30) (13) (11) (42) (27)

- 6 16 30 23 3 - 1 5

(5) (15) (20) (16) (6) (2) (8)

5 23 3 2 21 - - 13 10

(6) (18) GB) dy) (5) (21) (16)

2 4 3 1 1 - - 3 1

(G3) (3) (3) (1) (1) (5) (2)

2 1 - 1 - ] 46 - 1

(2) (1) (1) () (61) (2)

28 24 54 38 32 1 20 10 14

G4) (19) (52) (25) (23) (2) (26) (16) (22)

2 " - = * 1 1 : F

(2) (2) (tr.)

11 63 104 - 45 2 6 28 35

(13) (50) (100) (32) (4) (1) (45) (55)

51 92 32 21 72 5 - 50 42

(62) (73) (31) (14) (51) (1) (81) (66)

8 15 1 1 1 1 - 15 -

(10) (12) () (1) (1) (2) (24)

19, 14 55 48 38 2 - 3 11

Q1) qd) (53) (32) 27) (4) (5) (17)

22 H. A. MARTIN & R. L. SPECHT

Species Life Group 4 Group 6 Group 7 Group 2 Group 9

form 4(8) 4(9) 610) 611) 712) 713) 2, 9114) 915)

Haloragaceae

Gonocarpus tetragynus Ch 17 ) 1 1 3 9 3 3 2

(21) (4) (1) (1) (2) (19) (tr.) (5) (3)

G. humilis Ch 21 - - 7 - a 26 a r

(26) (34)

Mimosaceae

Acacia continua N 5 - - - 7 = bs a -

(6)

A, melanoxylon MM - 8 - 1 - 3 - 5 3

(6) (1) (6) (8) (5)

A. myrtifolia N 82 126 - - - 1 62 62 63

(100) (100) (2) (82) (100) (98)

A. pycnantha M 15 28 21 43 56 27 60 12 16

(18) (22) (20) (29) (40) (58) (79) (19) (25)

A. retinodes M 1 2 - 1 1 - - 2 -

(1) (2) (1) (1) (3)

A. rotundifolia N - 6 16 10 5 1 6 1 5

(5) (15) (7) (4) (2) (1) (2) (8)

A. rupicola N - - - - - 2 6 - -

(4) (1)

A, spinescens N 2 - - 1 - 1 50 ~ -

(2) (1) (2) (66)

Myrtaceae

Calytrix tetragona N 30 11 1 12 1 5 48 8 3

(37) (9) (1) (8) (1) (11) (63) (13) (5)

Leptospermum myrsinoides N 70 89 104 150 - - 54 57 32

(85) (71) (100) = (100) (71) (92) (50)

L. prickly (syn.

L. juniperinum) N 2. 4 5 3 6 2 5 3 1

(2) (3) (5) (2) (4) (4) (1) (5) (2)

Pittosporaceae

Bursaria spinosa M 3 4 - 2 6 2: - - 4

(4) G3) (1) (4) (4) (6)

Cheiranthera alternifolia N 9 - - - 2 I - - -

(11) () (2)

Poaceae

Danthonia sp. &

Stipa sp. H 9 15 - 5 16 14 31 6 9

(1) (12) G3) (1) (30) (41) (10) (14)

Proteaceae

Adenanthos terminalis N 2 2 - 1 - - 26 - 2

(2) (2) (1) (34) (3)

Banksia marginata M 22 23 1 5 > 6 5 18 B)

(27) (18) (1) (3) (4) (13) (1) (29) (8)

B. ornata N 2 2 - 1 - - 10 - 2

(2) (2) (1) (13) (3)

Conospermum patens N - - - 1 - - 3 7 hs

(1) (tr.)

Grevillea lavandulacea N 22 14 3 9 6 2, 18 9 5

(27) (1) (2) (6) (4) (4) (24) (15) (8)

Hakea rostrata N 65 58 78 111 29 3 26 42 16

(79) (46) (15) (74) 2) (6) G4) (68) (25)

H. rugosa N 1 1 2 5 3 - 26 - 1

(1) () (2) G) (2) G4) (2)

H. ulicina N 28 32 3 8 - 3 41 27 5

(34) (25) G) (5) (6) (54) (44) (8)

SCLEROPHYLL VEGETATION IN THE MOUNT LOFTY RANGES 23

Species Life Group 4 Group 6 Group 7 Group 2 Group 9

form 4(8) 4(9) 6(10) 6(11) = 7112) 7(13) 2 914) 9(15)

Isopogon ceratophyllus — N 25 13 - 5 - 1 21 13 -

(30) (10) (3) (2) (28) (21)

Persoonia juniperina N 3 7 - 1 - - - 6 l

(4) (6) (1) (10) (2)

Rhamnaceae

Cryptandra tomentosa N 4 - - - - - 11 - -

(5) (14)

Spyridium parvifolium N 1 7 15 20 8 1 - 3 4

(1) (6) (14) (13) (6) (2) (5) (6)

S. spathulatum N 3 1 - - - 3 - - 1

(4) (1) (6) (2)

S. vexilliferum N 12 6 - - - 1 - 5 1

(15) (5) (2) (8) (2)

Rubiaceae

Correa reflexa N 2 - - 1 - - 21 - -

(2) (1) (28)

Santalaceae

Exocarpos cupressiformis M 13 26 2 4 9 5 4 17 a

(16) (21) (2) (3) (6) (1) (1) (28) (14)

Sapindaceae

Dodonaea viscosa N 6 3 - 1 1 8 - 1 2

(7) (2) () d) (17) (2) (3)

Scrophulariaceae

Prostanthera behriana N - - - - - - 8 - -

(11)

Stackhousiaceae

Stackhousia monogyna = Ch 15 2 - 1 - - - 2 -

(18) (2) (1) (3)

Thymelaeaceae

Pimelea flava

subsp. dichotoma N - l - - 1 - 14 - 1

(1) (1) (18) (2)

P. glauca Ch - 1 - - - - 22 - 1

(1) (29) (2)

P linifolia N 20 30 44 52 35 6 - 7 23

(24) (24) (42) (35) (25) (13) (11) (36)

P. octophylla N 2 - - - - - 8 - -

(2) (11)

Tremandraceae

Tetratheca pilosa N 51 6l 46 40 55 2 - 29 32

(62) (48) (44) (27) (39) (4) (47) (50)

Xanthorrhoeaceae

Lomandra fibrata H 7 44 27 31 34 1 - 25 19

(9) (35) (26) (21) (24) (2) (40) (30)

Xanthorrhoea quadrangulata) M 11 2 - 1 - 8 - - 2

(13) (2) (1) (17) (3)

X. semiplana N 62 46 57 79 35 5 68 24 22

(76) (37) (55) (53) (25) (21) (89) (39) (34)

Transect sites

Inglewood 1 33 102 130 97 18 - - 33

x=414 (2) (26) (98) (87) (69) (38) (52)

Morialta 60 15 - 6 6 15 - 10 5

x=117 (73) (12) (4) (4) (32) (16) (8)

H. A. MARTIN & R. L. SPECHT

Species

Life

form

Waterfall Gully

x= 141

Belair N.P.

x= 89

Blewitt Springs

x=91

Total transects

LU= 852

Group 4 Group 6 Group 7 Group 2 Group 9

4(8) 4(9) 6110) 6(11) 7012) 713) 2 9114) — 9(15)

11 55 2 7 7 4 - 46 9

(13) (44) Q) (4) (5) (9) (74) (14)

6 20 - 6 31 6 - 6 14

(7) (16) (4) (22) (12) (10) (22)

4 3 - 1 - 4 76 - 3

(5) (2) (1) (9) (100) (4)

82 126 104 150 141 47 76 62 64

(100) (100) (100) (100) (100) (100) (100) (100) = (100)

Transactions of the Royal Society of S. Aust. (2005), 129(1), 25-38.

MACRO-INVERTEBRATES CAPTURED IN ARTIFICIAL SUBSTRATES IN THE

RESTORED WATERVALLEY WETLANDS IN SOUTH AUSTRALIA.

by J. M. WuiTe* & T. C. R. WHITE*

Summary

Wuire, J. M. & Waite, T. C. R. (2005). Macro-invertebrates captured in artificial substrates in the restored

Watervalley Wetlands in South Australia. 7rans. R. Soc. S. Aust. 129(1), 25-38, 31 May, 2005.

Ninety seven species of macro-invertebrates were collected from six wetlands in the upper south east of South

Australia between 1992 and 1995, During the study the salinity of the wetlands fluctuated between fresh or

slightly saline and moderately saline; in two instances the salinity temporarily exceeded that of sea water. These

changes in salinity had no apparent negative affect on the abundance of any group of macro-invertebrates with

the possible exception of Trichoptera in one wetland. This paper provides basic data about these wetlands before

the start of a major drainage scheme currently under construction. The information gathered here will improve

the active management of these wetlands when the scheme is completed.

Key Worps: Macro-invertebrates, wetlands, fluctuating salinity, tolerance to salinity.

Introduction

The Watervalley Wetlands are a series of alkaline,

shallow and fresh or slightly saline to moderately

saline, temporary, semi-permanent or permanent

wetlands in the upper south east of South Australia

(White 1999a). They were hydrologically restored

between 1984 and 1991. They are mostly connected

either by natural broad, shallow watercourses or by

drains. In historic times these wetlands were

connected to the ocean via Salt Creek and the

Coorong (Fig. 1) but this connection was disrupted

by drains further to the south. The Watervalley

Wetlands are of national conservation significance

(Jaensch & Auricht 1989; Nicholson 1993) and

qualify for listing as Wetlands of International

Importance under the Ramsar Convention although

they have not yet been so listed. The wetlands, at the

time of the study, were fed mainly by fresh to very

slightly saline water which flows through the system

of drains and watercourses from the south-east

towards the north-west in all but extremely dry years.

The median annual rainfall at Naracoorte, the nearest

weather station upstream of the wetlands,

is 577 mm. Rainfall decreases sharply towards

the north and is 456 mm at Keith, the nearest

station downstream (Commonwealth Bureau of

Meteorology). As the gradient of the watercourses is

of the order of only 1:5000, except at a few isolated

points, the rate of flow is extremely slow. The

wetlands vary in size from 100 hectares (Jip Jip) to

2700 hectares (Mandina Marshes) and are managed

“School of Natural and Built Environments, Mawson Lakes

Campus, University of South Australia, Mawson Lakes, SA 5095.

* School of Agriculture and Wine, Waite Agricultural Research

Institute, University of Adelaide, Glen Osmond, SA 5064.

primarily for conservation. White & Brake (1995)

described them and outlined their history. The

chemistry of their waters, particularly with respect to

salinity, was discussed by White & Brake (1995) and

White (1999a). Six of the wetlands (Jip Jip, Mandina

Marshes, Mandina Lakes, Cortina Lake, Bonneys

Camp South and Bonneys Camp North) are

discussed in this paper.

All of the Watervalley Wetlands are actively

managed with the aim of maintaining or enhancing

their value as habitat for native wildlife, particularly

native and migratory waterbirds. A program of

monitoring water chemistry and various biological

attributes of the wetlands commenced in April 1992.

The purpose was to observe succession in these

newly rehabilitated or restored wetlands and to

monitor their ongoing “health” in a region that is

subject to rising groundwater and increasing dryland

salinity (Clark ef a/. 1991). This dual problem is

currently being addressed by a drainage scheme

to lower the ground water: the Upper South

East Dryland Salinity and Flood Management

Programme. As a result of this scheme the wetlands

are currently (2004) receiving more saline water

from the new groundwater drains than they received

previously and, overall, salinity has increased and the

periodical influx of freshwater has diminished. When

the scheme is completed it should allow for the

restoration of the natural late winter and spring

inflow of fresher water.

The importance of invertebrates in the diet of

waterbirds is well documented (e.g. Hill et al. 1986;

Cooper & Anderson 1996; Cox et al. 1998), This

paper discusses the results of monitoring macro-

invertebrates between 1992 and 1995. The results

provide baseline data for any future monitoring of

the effects of the drainage scheme and its impact,

J.M. WHITE & T. C. R. WHITE

c i

ah Us

‘

.» Pretty

_Lagoon —

Seaors he

Johnnys:

SOUTH AUSTR

“2 t

me \ Jip Jip

Naracoorte

Skm East —> |

440° 00

Fig. 1. The upper south east of South Australia showing the Watervalley Wetlands.

either positive or negative, on the native animals and

plants of the wetlands.

Methods

Invertebrates were collected at intervals of four to

six weeks between August 1992 and the late spring

or early summer of 1994. Further collections were

made in 1995.

One invertebrate sampling site was established at

each of the six wetlands between April and July

1992. Five artificial substrates were used at each site.

They were suspended from star pickets placed five

metres apart along a transect at an angle of 90° to the

shore commencing where the water was 60 cm deep

at the time the transect was established. This gave a

range of potential depths of up to one metre along a

transect. At Bonneys Camp South this configuration

was not feasible as the lagoon shelves very steeply to

a depth of three to four metres. The transect at this

site ran parallel to the shore at a depth of 60 cms

when first established.

The artificial substrates were similar to those used

by Bennison ef al. (1989) and made of plastic mesh

Gutterguard™ baskets containing one and a half

onion bags made of red, loosely woven nylon mesh

with a total surface area of 4256 cm’. The baskets

were each 17 cm high with a 16 cm diameter circular

MACRO-INVERTEBRATES IN RESTORED WETLANDS 27

base. Two small plastic vials filled with sand were

placed in the bottom of each basket to act as ballast.

The substrates were suspended from the star pickets

by nylon rope so that they rested on the bottom of the

wetland. They were left undisturbed for four to six

weeks. The macro-invertebrates were then collected

by placing a dip net under each substrate with as little

disturbance as possible and bringing it to the surface.

The contents of the substrate were then washed into

the dip net and concentrated in a vial attached to the

base of the net. Small fish and yabbies (Cherax

destructor) were released. Samples were preserved

in 70% ethanol on site.

At the same time as the invertebrates were

collected the depth of water was recorded and later

converted to the Water Level Index of Tamasier &

Grillas (1994). Conductivity, turbidity and major

chemicals in the water were analysed (reported in

White & Brake 1995). Conductivity is an indirect

measure of salinity and, in the text where

conductivity measured in mS/cm is converted to

salinity in g/L, the conversion is g/L = 0.68 x

conductivity in mS/cm (Hart et al. 1991). In this

paper the term “saline” is used to describe water with

conductivity greater than 4.4 mS/cm (3 g/L). Water

with conductivity of 4.4 mS/cm to 14 mS/cm is

described as being slightly saline and with

conductivity 14 mS/cm to 50 mS/cm as moderately

saline. From August 1993, the dominant species and

total cover of submerged plants, as a percentage,

were recorded in a 50cm quadrat surrounding each

point. Percentage cover of vegetation and depth of

water was then reduced to a mean for each transect.

In the laboratory each invertebrate sample was

tipped into a glass Petri dish, 20 cm in diameter,

resting on a sheet of paper. Lines had been ruled on

the paper to partition the area of the bottom of the

dish into eight equal “pie” segments. The sample was

stirred and allowed to settle as evenly as possible

across the whole dish.

Counting was done under a x 4 dissecting glass as

follows:

* All animals in the dish: large animals such as

adult coleopterans, late instar odonates, spiders,

and the larval cases of trichopterans (only the

cases of trichopterans were counted as some, but

not all, larvae left their cases when placed in

alcohol).

¢ All animals in the right hand half of the dish:

earlier instars of the above when they were

numerous, and small snails.

* All animals in the top right hand quarter of the

dish: small and numerous early instars.

* All animals in the top right hand eighth of the

dish: very small and very numerous amphipods,

larval chironomids and ostracods.

For purposes of comparison and calculation all

counts were standardised at one-cighth-equivalents

of the total sample. Counted animals were placed in

a separate vial and the residue of the sample returned

to the vial in which it had originally been placed in

the field. Identified specimens from each site are

lodged with the Museum of South Australia.

Statistical calculations were made using Pop

Tools™ version 2.5.8 (Hood, 2003) and Microsoft

Excel™,

Results

Bonneys Camp North and Bonneys Camp South

held water throughout the sampling period. The sites

at Cortina Lakes and Mandina Lakes dried briefly in

the autumn of 1994, although other areas of Cortina

Lakes continued to be inundated. Except for the

spring of 1994, all but several deeper waterholes

downstream of the sampling area at Mandina

Marshes, were dry from January 1994 to July 1995.

These periods of drying were due to low rainfall. Jip

Jip was drained for management purposes in the

early winter of 1993 and it did not begin to fill again

until September 1995. The water level at all sites

fluctuated seasonally. The number of times that each

site could be sampled was dictated by water levels.

Bonneys Camp North was sampled 21 times;

Bonneys Camp South 22; Cortina Lakes 23; Jip Jip

seven; Mandina Lakes 16 and Mandina Marshes 21.

Full details of the water chemistry at each site

during the study were published in White & Brake

(1995): a summary of the range and median value of

conductivity, nitrates and orthophosphate is given in

Table 1. Water levels were published in White

(1999a). Conductivity at all sites varied during the

TABLE |. Range (and median value) of certain chemical attributes from June 1992 to December 1995.

Bonneys

Camp North

Bonneys

Camp South

Cortina

Lakes Lake

Mandina Mandina

Marshes

Jip Jip

Conductivity mS/em —4,30-16.30 (7.61) 4.2-9.59 (6.45)

Nitrate mg/L 0.2-2.0 (0.8) 0, 1-2.6 (0.7)

Orthophosphate mg/L

3.32-69.30 (8.91)

<0,1-2.6 (0.7)

0.01 - 0.88 (0.04) <0.10 - 0.40 (0.05) 0.01 - 0.83 (0.04) 0.01 - 0.76 (0.15)

1.30-5.58 (2,26) —2.37-116.00 (14.63) 1.60-16.80(5.91)

<0.1-5.3 (0.4) <0.1 - 3.7 (0.6) <0.1 - 0.6 (0,2)

<0.01 - 0.94 (0.60) <0.01 - 0.65 (0.05)

study (Table 1; Appendix) with Cortina Lakes and

Mandina Lakes each having a spike of extremely

high conductivity in March 1994 as water levels

receded. The water at all sites, except Jip Jip, was

slightly saline for most of the time (conductivity < 14

mS/cm or salinity < 10 g/L). Jip Jip was fresh to very

slightly saline. The spikes of high salinity in both

Cortina and Mandina Lakes, which occurred in

March 1994, exceeded the salinity of sea water (sea

water = 35g/L or approximately 50 mS/cm).

Bonneys Camp North, Cortina Lakes and Mandina

Lakes were well vegetated but Mandina Marshes and

Bonneys Camp South were sparsely vegetated in the

earlier part of the study. Jip Jip was sparsely

vegetated throughout. The hydrology of the two

Bonneys Camp wetlands was restored in 1991 and, in

the first year of the study, little aquatic vegetation

had developed on the bare sandy bottom of Bonneys

Camp South but the sedgeland which covers the bed

of Bonneys Camp North was apparently unaffected

by inundation. This sedgeland was relatively dense

(mean cover of all species combined = 50%) and

dominated by Baumea juncea and Hypolaena

fastigata. In contrast Bonneys Camp South was

devoid of vegetation until December 1993 when a

submerged community dominated by Lepilaena spp

and charophytes (mean combined cover = 25%)

developed. From the beginning of the study until

November 1994 the sampling area at Cortina Lakes

TABLE 2. List of species of invertebrates recorded at each site.

J.M. WHITE & T. C. R. WHITE

was dominated by emergent Baumea arthrophylla

and submerged charophytes with a mean combined

cover of 65%. Between that time and the next

sampling in November 1995 the water level receded

below the sampling site for over six months and the

sedges and charophytes were replaced by Ruppia

megacarpa when the water level rose again. The

sampling site at Mandina Lakes was dominated by a

mix of Ruppia megacarpa and Lepilaena spp with a

mean cover of 65%. The sampling site at Mandina

Marshes had very sparse vegetation until it reflooded

with fresh water after the drought and the site was

colonised by a mix of Potomogeton pectinatus and

P. crispus.

Ninety seven species of macro-invertebrates were

collected during the study (Table 2) and of these, 25

species were found at only one site. Fifteen species

were found at two sites, 15 at three sites, seven at

four sites, 11 at five sites and 20 were common to all

sites. Four species could not be assigned to a specific

site because they had been identified from

preliminary general samples and were not found

again. Cortina Lakes (59 species), Mandina Lakes

(56 species) and Mandina Marshes (54 species) were

the richest in species, Bonneys Camp North (51

species) and South (49 species) were slightly less so,

and Jip Jip (37) had fewest.

The overall mean number of each of the major

taxonomic groups of invertebrates at each site is

Taxa BCN BCS

CL JJ ML MM

Cnidaria (hydra)

Clavidae

Cordylophora sp

Hydra indet

Total species of 2

Cnidaria at each site

Mollusca

Gastropoda( snails)

Planorbidae

Glyptophysa tenuilirata

Hydrobiidae indet

Helicarionidae

Echonitor nr cyrtochilus

Total species of l 1

Gastropoda at each site

Hirudinea (leeches)

Glossiphonidae

Glossiphonia sp (site not specified)

Hirudinidae

Bassioanobdella sp (site not specified)

Total number of species = 2

Crustacea

Ostracoda: seed shrimps

Cyprinidae (indet)

eNO ONS

MACRO-INVERTEBRATES IN RESTORED WETLANDS

TABLE 2. List of species of invertebrates recorded at each site cont.

Taxa BCN BCS

CL JJ

Mytilocypris sp v

Notodromatidae (indet) v ov

Total species of Ostracoda 2 2

Amphipoda (scuds)

Ceinidae

Austrochiltonia australis v v

Austrochiltonia subtenuis

Crangonyctidae (indet) site not specified

Eusiridae (indet) v

Total n. species 1 1 2 3

of Amphipoda

Decapoda (freshwater crayfish)

Parastacidae

Cherax destructor v v v v

Total species of Decapoda 1 1 1 1

Arachnida:

Acarina (mites)

Arrenuridae

Arrenurus sp | v vo v v

Arrenurus sp 2

Limnocharidae

Rhyncholimnochares sp v v v v

Eylaidae

Eylais sp v

Hydrachnidae

Hydrachna sp v

Pionidae

Piona sp v

Total species Acarina 2, 2 5

Araneae( spiders)

Lycosidae

Trochosa tristicula phegea v

Trichosa sp v

Tetragnathidae

Tetragnatha sp v

Linyphiidae

Erigona prominens v v

Total species Araneae l I 3

Insecta

Ephemeroptera (mayflies)

Baetidae

Cloeon sp

Total species Ephemeroptera

Odonata: Zygoptera (damselflies)

Coenagrionidae

Coenagrionidae juv. v

Austroagrion coeruleum Jv Jv

Austroagrion watsoni

Ischnura heterosticta v

Ischnura sp

Xanthagrion erythroneurum v

Lestidae

Austrolestes annulosus v

Austrolestes analis v

Austrolestes juv v

So ONESCNEN

—

J. M. WHITE & T. C. R. WHITE

TABLE 2. List of species of invertebrates recorded at each site cont.

Taxa

BCN

BCS

Megapodagrionidae

Argiolestes icteromelas (site not specified)

Total species Zygoptera

Odonata: Anistoptera (dragonflies)

Aeschnidae

Aeschna brevistyla

Hemianax papuensis

Hemicorduliidae

Hemicordulia tau

Libellulidae

Austrothemis nigrescens

Diplacodes bipunctata

Diplacodes haematodes

Total species Anisoptera

Total species Odonata

Hemiptera (bugs)

Corixidae

Agraptocorixa eurynome

Agraptocorixa sp

Diaprepocoris barycephala

Micronecta sp

Sigara australis

Sigara sp

Naucoridae

Naucoris congrex

Notonectidae

Anisops sp

Anisops thienemanni

Total species Hemiptera

Coleoptera ( beetles)

Dytiscidae

Platynectes decempunctatus

Allodessus bistrigatus

Antiporus femoralis

Antiporus gilberti

Hyphydrus elegans

Megaporus gardeneri

Megaporus hamatus

Megaporus larva

Necterosoma penicillatum

Sternopriscus tasmanicus

Rhantus suturalis

Lancetes lanceolatus

Hydrophilidae

Limnoxenus zealandicus

Enochrus eyrensis

Paracymus pygmaeus

Stenophilus marginicollis

Berosus majusculus

Berosus discolor

Berosus veronica

Berosus larva

Halplidae

Haliplus sp

Total species Coleoptera

Ss¥vNNN

VNNNN

NNN

\N\N

SONGN

Sen

\N NNN

SNe SONTREN ON NN

NNN

ANS

MACRO-INVERTEBRATES IN RESTORED WETLANDS 31

TABLE 2. List of species of invertebrates recorded at each site cont.

Taxa BCN BCS CL JJ ML MM

Diptera (flies)

Ceratopogonidae v

Nilobezzia sp v v

Chironomidae

Ablabesmyia sp

Chironomus sp v v

SoS

Cladotanytarsus sp

Cladopelma sp

Cricotopus sp

Dicrotendipes sp

Larsia sp

Sani SON

SENSORS

Parachironomus sp

Paramerina sp

SO NENEN

NANA SS

SNES

Polypedilum nubifer

Polypedilum sp

Procladius sp

Tanytarsus spp

Chironominae indet

Ephydridae indet

Stratiomyidae indet

Tabanidae indet

Total species Diptera

Trichoptera (caddis flies)

Ecnomidae

Ecnomus cygnitus v v v

Ecnomus sp v v

Hygrocloatidae indet

Leptoceridae

Notalina salina

Notalina spira

Oecetis spp

Symphitoneuria opposita

Triplectides australis v v v v

Total species Trichoptera 5 6 5 3

Lepidoptera (moths)

Pyralidae

Nymphulinae indet v

Nymphalinae sp 40!

Total species Lepidoptera 1

Total number of species 51 49 59 37 54

BCN = Bonneys Camp North; BSC = Bonneys Camp South; CL = Cortina Lakes; JJ = Jip Jip; ML = Mandina Lakes;

MM = Mandina Marshes. |= sp 40 in SA Water Reference Collection.

SSN a) QVNNNNNNRNNNRS

By, ANY \o \QN\N

SONA = SNS

N oo \N

SSN Bm WON ONSEN Se NoSNG SNe

BSS SREN SN =a NN ONE \

IRENE

NyQAN

TABLE 3. Overall mean number (and standard deviation) of major invertebrate groups at all sites.

Taxa BCN BCS CL JJ ML MM

Amphipoda 50.19 (81.2) 54.9 (36.1) 113.4 (97.6) 410.4 (355.4) — 188.3 (303.9) 583.8 (555.5)

Ostracoda 36.57 (32.5) 29.9 (41.4) 12.0 (20.1) 4.75 (5.4) 45.9 (31.9) 2.5 (3.1)

Trichoptera 7.24 (7.4) 7.9 (6.4) L121) 28.0 (24.0) 9.3 (5.0) 6.8 (6.0)

Zygoptera 3.04 (3.8) 4.8 (3.4) 2.8 (2.1) 7.7 (5.95) 3.4 (4.5) 7.6 (6.7)

Anisoptera 0.2 (0.27) 0.2 (0.3) 0.4 (0.3) 0.3 (0.3) 0.3 (0.1) 1.3 (3.0)

Diptera 36.57 (21.45) 27.4 (23.2) 10.1 (10.0) 12.4 (4.2) 24.9 (48.4) 22.6 (17.8)

Coleoptera 3.22 (1.8) 1.7 (1.8) 0.2 (0.2) 1.7 (1.4) 4.6 (4.8) 0.3 (0.3)

Hemiptera 0.06 (0.08) 0.1 (0.1) - 1.0 (1.6) - -

Acarina 9.44 (10.95) 2.4 (7.8) 0.4 (0.4) 0.55 (0.7) 0.9 (0.7) 6.7 (11.2)

Gastropoda - - 0.6 (1.6) 10.9 (9.8) 0.2 (0.3) 10.9 (3.7)

32 J, M. WHITE & T. C. R. WHITE

given in Table 3. Histograms showing the mean

number for each sampling date, together with

conductivity at the time, can be found in the

appendix. Cortina Lakes was the only site where

enough samples were collected both before and after

the spike of extremely high conductivity (69.3

mS/cm) to get a statistically valid indication of the

effects on macro-invertebrates of such relatively

extreme conditions. Student’s ¢ test for the difference

between these two sets of samples showed a

significant increase in abundance of Ostracoda,

Odonata and Acarina and a significant decrease in

Trichoptera. There was no significant difference in

the abundance of Amphipoda, Diptera, or Coleoptera

(Table 4).

Discussion

All of the wetlands except Jip Jip are part of the

Bakers Range Watercourse and are connected in

times of high flow (Fig. 1). When Jip Jip (in the

Marcollat Watercourse, several kilometres to the east

of the Bakers Range Watercourse) overflows, water

from it passes through a drain into the Bakers Range

Watercourse and thence to the other wetlands studied

(White & Brake 1995). This provides for a degree

of migration by flightless macro-invertebrates

throughout the system.

The wetlands of the Upper South East are

dependent on fresh water flowing north west from

the upper reaches of the catchment which extends

TABLE 4. Results of Student’ ¢ test comparing mean

numbers of major invertebrate groups after spike in

salinity in March 1994 at Cortina Lakes.

A93-M94 Ap94-N94

Amphipoda Mean=104.6 Mean=130.6

Var=7104.8 Var=3071.2

P=0.207

Ostracoda Mean=14.9 Mean=73.7

Var=373.4 Var= 1981.6

P =0.010

Trichoptera Mean=0.8 Mean=0.2

Var=0.6 Var=0,02

P=0,.006

Odonata Mean=6.0 ean=13,2

Var=44.6 Var=24.7

P=0.008

Diptera Mean=16.0 Mean=8.1

Var=209.4 Var=67.2

P=0.063

Coleoptera Mean=0.5 Mean=1.1

Var=0.57 Var=1.31

P=0,147

Acarina Mean=0.7 Mean=1 1.2

Var=0.61 Var=27.0

P=0.002

into western Victoria. Local run-off provides some

inflow in winter although the amount is probably

much less important. The relationship between the

wetlands and groundwater is unknown. The very

slight gradient to the north west means that the flow

of water is slow and evaporation rates are high,

accounting for the higher salinity in the more

northerly wetlands. Mandina Lakes and Cortina

Lakes are fed only by water entering through

controlled inlets from Mandina Marshes. Neither of

these wetlands has an outlet, therefore as water levels

recede, their salinities are relatively higher than the

other wetlands. The natural flow throughout the

system has been disturbed by drainage but the

current scheme provides for both surface water and

drainage water to be managed to benefit the wetlands

(NRCSA 1993). This should allow the hydrological

regime of the wetlands before drainage to be re-

established, thus potentially conserving the habitat

for species currently found in the wetlands.

Mandina Marshes and Cortina Lakes are the least

disturbed of all the sites. Even after the overland flow

to the more northerly wetlands was reduced by 50%

in 1964 because of drainage (EWS 1991), the deeper

pools of both these wetlands retained water

permanently (T. K. Brinkworth pers.com.). Both the

riparian and the aquatic vegetation of these wetlands

appear to support this notion (J. M. W. unpublished

data) and both areas have surrounding terrestrial

vegetation that has not been cleared. This relative

stability is consistent with these two bodies of water

having a greater number of species than most of the

others, although it does not seem to explain the high

number of species on Mandina Lakes. The whole of

this wetland was grazed pasture from the 1960s until

its restoration in the late 1980s (T. K. Brinkworth

pers. com.). Jip Jip was only sampled seven times

because it was drained in the autumn of 1993 and it

did not receive any water until the spring of 1995.

This necessarily reduced sampling effort could

account for the paucity of species there; although the

fact that its hydrology was disturbed both prior to

and during the study could also be a factor.

There was no apparent negative relationship

between salinity (measured as conductivity) and the

abundance of any groups of macro-invertebrates

found at any of the study sites except, possibly, for

Trichoptera at Cortina Lakes. It is frequently stated

that increases in salinity lead to a decline in the

number of species (e. g. Hart ef a/. 1991; Nielsen et

al. 2003) and saline wetlands are therefore

sometimes regarded as being “less important” than

freshwater systems. However, this decrease in

species occurs at levels of salinity generally below |

g/L, below any yet measured in the Watervalley

Wetlands cited here except rarely and briefly in Jip

Jip. Furthermore, while there may be fewer species

MACRO-INVERTEBRATES IN RESTORED WETLANDS 33

in more saline water, they can be much more

abundant (Kingsford & Porter 1994). This could

make saline wetlands much more important as a

source of food for waterbirds.

In this study, however, the greatest number of

species was found in the two most saline lakes:

Cortina (59) and Mandina (56). Mandina Marshes,

with 54 species also had a relatively rich fauna. The

freshest wetland, Jip Jip, had fewest species but

because it was sampled less frequently than the

others it would be unwise to make too much of this

observation. The methods of collection used in this

study trap only those species which will colonise

substrates and micro-crustaceans were not counted,

therefore our species lists do not represent the true

number of species present in any of the wetlands.

At Cortina Lakes there was a. statistically

significant increase in the abundance of Ostracoda,

Odonata, and Acarina after the spike in salinity

(measured as conductivity) in autumn 1994 whereas

the abundance of Trichoptera decreased after that

event. There was a twofold increase in the abundance

of Coleoptera but the variance was such that it was

not statistically significant. However, caution is

advised in attributing these changes to the sharp rise

in salinity. The increase in abundance in both

Odonata and Coleoptera began before the event and

there was a very long lag time for Ostracoda and

Acarina. It is possible, however, that the decline in

the abundance of Trichoptera was due to higher

salinity. This has been reported elsewhere (Hart ef al.

1991), but Kefford et al. (2003) found that Notalina

spira and species of the genus Trip/ectides other than

those found in Cortina were tolerant of conductivity

greater than 25 mS/cm. So, even for this group, it is

not certain that the observed changes can be

attributed to changes in salinity.

This study was devised to find out whether there

were discernable trends in the abundance of macro-

invertebrates following restoration, not to measure

the effects of increases in salinity on individual

species or orders. Further studies are needed to

ascertain the tolerance of the species present to the

sometimes quite large seasonal increases in salinity

because the hydrology, in regards to quantity, quality

and timing of flows, of these wetlands will be

actively managed through the Upper South East

Dryland Salinity and Flood Management Scheme.

The tolerance of some of the species present is

known (see Hart ef al. 1991; Kefford et al. 2003) and

it seems, not unexpectedly, that these macro-

invertebrates are able to cope with such widely

fluctuating salinities. However, it is also known that

the tolerance to increases in salinity varies at

different stages of the life cycle (Hart et a/. 1991),

Therefore, it does not follow that all species present

could tolerate constant salinities around the median

levels observed during the study. Nielsen ef al.

(2003) found that hatching of zooplankton and

germination of plants was affected by increases in

the salinity of fresh water. Similar effects could be

present in these systems whose salinity can fluctuate

widely within a single year and it may be that the late

winter-spring inflow of less saline water is essential

for reproduction of some species. This, too, requires

further study.

Our results show that the Watervalley Wetlands

experienced large seasonal fluctuations in salinity

and supported a diverse macro-invertebrate fauna

before the current drainage scheme commenced. It

would be prudent, therefore, to manage water

regimes in the Watervalley Wetlands so that these

seasonal fluctuations in salinity are maintained

unless it can be demonstrated that these slightly to

moderately saline systems can maintain their mix of

species of both plants and animals without seasonal

periods of lower salinity for successful recruitment.

Acknowledgements

Travel and collection of field data for this paper

were supported by grants from the University of

South Australia and the Wildlife Conservation Fund

of South Australia.

We would like to thank the following people for

their assistance in this project: Lynn Brake

(University of South Australia) who helped devise

and set up the monitoring program and assisted with

initial field work; many students of Conservation

and Park Management (University of South

Australia) who assisted with field work; Tracy

Corbin, Chris Madden and Paul McEvoy (Australian

Water Quality Centre), Chris Watts and John

Bradbury (SA Museum), and John Hawking

(Murray-Darling Freshwater Research Centre,

Albury) who identified specimens and Dr Roger

Clay (University of South Australia) for advice on

statistics. David Britten (University of South

Australia) prepared the base map used for this paper.

We are also grateful to Pat and Tom Brinkworth of

Ninga Ninga Station, Kingston SE for their

hospitality during field trips and for allowing us

unfettered access through their property to the

wetlands.

34 J. M. WHITE & T. C. R. WHITE

References

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Macro-invertebrates of the River Murray. Murray-

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CLarRK, R., Kotwicki, V. & TOMLINSON, G. (1991).

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Cooper, C. B. & ANDERSON, S. H. (1996). Significance of

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the internet. URL http://www.cse.csiro.au/poptools.

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Relative salinity tolerance of macro-invertebrates from

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KINGSFORD, R. T. & Porter, J. L. (1994). Waterbirds on an

adjacent freshwater lake and salt lake in arid Australia.

Biological Conservation. 69: 219-228.

NICHOLSON, C. (1993). Conservation parks and wetlands in

the upper south east — surface water recommendations

and an assessment of impact associated with rising

groundwater. Department of Environment and Land

Management South Australia.

NIELSEN D. I., Brock, M. A., CROSSLE, K., HARRIS K.,

HEALEY, M. & JARosINskI, I. (2003). The effects of

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hatching from two wetland sediments. Freshwat. Biol.

48: 2214-2223.

NRCSA (Natural Resources Council of South Australia).

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Management Plan. Department of Environment and

Natural Resources SA.

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community. Biol. Conserv. 70: 39-47.

Wuirte, J. M. (1999a). Seasonal variation in salinity in the

Watervalley Wetlands in the south east of South

Australia. Trans R. Soc. S. Aust. 123 (2), 77-80.

(1999b). Watervalley Wetlands and Heritage

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University of South Australia for Wetlands and Wildlife,

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& BRAKE, L. A. (1995). Description, history,

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15 (3), 247-257.

MACRO-INVERTEBRATES IN RESTORED WETLANDS

APPENDIX. Mean number of macro-invertebrates and conductivity (mS/cm) at each wetland.

Amphipoda

Bonneys Camp North | Bonneys Camp South

400 —_ 20 | 1000 — 20

350 beeeuecee (mE A mphipoda

| @ 300 |, —e—Conductivity 415 | yp 800 [=e Conductivity] ° 1 15

= 3 250 al 2% 600 €

§ 2 200 105 | & 8 105

| os a 2 400 s i age

| S & 150 “ =o ey go £

< 100 © 45 © 300 We 5

| he | |

| | |

| Aaalla a. ato o an, Me, afl ans, ie 5

POZO GL MSSSEVOULNE>SSEP= EE Z | BOZO TEES EPOOO TEPESEPZ zPZ

@ oO B 2 | oO ©

+ Date N Date a

Cortina Lakes Jip Jip |

400 — zu ____+ 80

350 + Saee 70 1200. {mmm A iphipoda] 7°

| = 300+ —e—Conductwity] gq | 1000 [—e—Conductvty| 15

z3 | a os

Z 8 250 50e| 2 800 foe

& = 200 +408 | es S|

2B 150 130 © ae S| ees

100 20 <

1 5

50 I] I 10 200 | al

° powoenessy TEEPE Aer Ee 0 MT | ee er ee Cae PEs

Peale an ¢ a = BOZO NESS POO SP ESSEZ FF

ue Date oi 8 Bate 8 {

Mandina Marshes | Mandina Lakes

2000 20 1000 80

] [aero ] | | {ummm Amp hipoda ]

| g 1800 + —#Conduernty 15 |g 800 |e conductivity +60 |

| 28 |, §) 2 8 600 = io 5 |

8 & 1000 [106] g 38 + 405

se k ” |. £| | 2 2 400 £

< 500, 5 2 20

li II. 200

om MiMer, wee, Sei eo o WIT eee, Lo

Fa POI a 4

& 1g woe OOo Se BE BOZUST ERE EM OURNE

i] Date “ | Ss al Date

Ostracoda

Bonneys Camp South | Bonneys Camp North

200 ——— 20 | | 140 + eT oe + 20

mmm Ostracoda ea mmm Ostracoda | |

150 —#— Conductivity 3 15 ie y [Re conductivity |

<8 a0:

22 60

=o |

S 404

20 |

| () : AL. - o |

| PUZOL TEES POOUL TEP ESEEZ zeez BOZOCNESES PNOULMEPESLPHZ ZY Z

oO 8 g ive} oO oe oonx< Cc o

| Date OY f s °° Tite &

Cortina Lakes Jip Jip

| 160 80 20 = ws 23)

| 140 | SMM Ostracoda 70 | | {mmm Ostracoda

| 5 g 120 {—e—conauetiity, @ || og 15 + |—e—conduetiity!| 45 |

3 iS 3 €]

§ 8 tag! | § 8 104 +109

2a 130E}) se |

9 ili | 20 1) 7 ls

() ol, ae, See ell lo

Zz | BOZO EES 2000 MP ESEEZ =zPze

| | a 8 & =

| s ‘Date a

Mandina Lakes Mandina Marshes

120 7 = __+ 80 a.

| mmm Ostracoda | 79 | noe Mmmm Ostracoda | nm

A 100 [Conductivity | 6g j 8 [=e—conducivty) 45

z 80 ae ae 225 a |

aren <8 6 §|

8 Sf, 7 106 |

= 8 4 a = a ” E

a I ELL Ie

o 1a Masteeto | 0 Me

Pa Jer ~_

POZOS NEES POOOS io WS PESSEZ

POZOSNEESPUOOL TE DESC EZ = DP

S Bate a

36 J.M. WHITE & T. C. R. WHITE

Trichoptera

va Pa — oy a

Bonneys Camp North Bonneys Camp South

_ 20 40 20

28 es Tiichopters T ‘mmm Trichoptera ]

20 {w= conductivity | 15 ge 30 {Tem Conduetivity ‘s +18

o z

A §| ls |

22 Ei |22 | |

BE =

~ Cortina Lakes | Jip Jip

6 80 a BD

[mmm Trichoptera [ | Trchoptera ||

s [—e=Conauctvty |. g @ 60 [—e—Conductivity|) 45

Ze 4] a Zo £

| $2 g 2 0+ |B

| =2 ol | ss | E

| ec “| | F 204 5

{ {

0 onal of aan flo

BOZO SESE (OMG SETS S<e= 5 z= PUZOS NEES POOOLTEPESEEZ EE Z

8 Bate & g 8 Bre. &

Mandina Lakes Mandina Marshes

20 + = 20

ee r Tachowtere] ithe ] mE Trichoptera

| —e—Conductivity || gq se 15 —e—Conduetivity | 45

z5 abe | — al 7 g i €

2 c a

§ 3 +40 3 | § 2 10 ” 10 a

= 8 | Elio = £ 7 5

a 20

| o?

é lo Id ) | rere ererarereraresneae | at,)

POZOCNEESPOOOLMEPESE PZ = PZ 1 | 2020, ESSEC SF SSKE= FE =

c © ox oo < IS go © ; 4 o oO a +h

@ 8 Cate 8 |] N Date

Odonata

a -_ 7 7. Re a i = "2a c =a ]

Bonneys Camp North Bonneys Camp South

_——_____— 20 16 _—____. 20

8 12 | —e—Conductivity 15 g ~e—Conductivity| |

S | |

3 10 3 2

o 8 Loge||

€

<« 6 Pat wo ©

8 aN, * +5 3g

= 21

nll, Elas i ++0 |

pozo nese SNOUCTEPESEPZ = % z POZOCMESP MOUS TE YESS EZ EER Z

i=] ied o< 7c 2 ry ase 2 c = sc. p< st 2 o< s

8 Date o ad Date a

= = ae ae ~ 1P as aa ==

| Cortina Lakes | Jip Jip

80 6 __7 20

@ gms Odonata | g 5 Odonata |

% 20 }|—e—Conductivity 60 @ [—e—conductivity 15

3 15 Bihuege 11.

9 40 3 S 3+ 10%

Z 10 & © 2+ ey

S 5 20 8 |

0 oO | O++ ttt ttt etl] 0

POZOCNZECHPWOUCNZSPESESSPZ FPS

BOZO NEES E000 WE PESSEZ 5 Pz a geese ‘2 ES<2 a €

8 Date a Ly Date

Mandina Lakes Mandina Marshes

| 16 7 =a 80, 25 ______,20

g { lm Odonata i me Odonata

@ 42 —e— Conductivity} EQ, 3 20 + —e— Conductivity! 45

£ 6 + €

8 8 + 40 a | 8 | 10

< | | € 10 | |

) 3 44 20 gs 5

o Mts meas ae Looe eeu seede a

FOZUS NESS FOOUK NE >ESSE= = 2 Zz BOOB TEESE MOOG" E”ES Sez g 2 z

Bg = Site & Cate

MACRO-INVERTEBRATES IN RESTORED WETLANDS

Diptera

Bonneys Camp North

12059" = ved 20

| [mmm Diptera i

D = — 15

a |

o {

“eal

Cortina Lakes

50 7 = im , 80

ummm Diptera |

p [=e=Conductivity | 60

& 30 §

z 40 3

< 20 Q

S 20 |

= 10

0 ttt o |

POZOSNEESPOOOS ME HESS EZ = Pa

aa Date a

200 + ; 80

FS | Lz |

@ 150 + | e—conduetivity 60

a +505

Zz 100+ + 40 4 3 |

c 30

gm, in|

| ois Mee re

it if

FO ZOOS NESS POCO TNS ES SEZ = ez 2 |

@ 8 g 8 |

Date

Coleoptera

Bonneys Camp North

a 20

gam Coleoptera

3 4 iberceee 415 |

z§ = ae =

| EZ <B950 5

| 88 9 0 9

zo ae ai

OR wer al

0 be aan dll _ —

EOZOG NESS POOL NEP ESE E= = 4 Zz

oa asa aa &

Mean N

Coleoptera

Mean N

Coleoptera

4 7 80

umm Coleoptera

3 | -e—Conductivity| + 60

40 Q |

20 |

bab O

POZO TEES PUOU TEP ESE EZ zPZz

é <e= Fe

rey 8 g 8

% Date

Mandina Lakes

145 —_.—+ 80

12 mmm Coleoptera | 7

40 —e—Conductivity

2 | 150 §

a © Q

i 30 £

4+ 20

2 Hee 10

0 ana ME to |

iz Zz

Date |

Cortina Lakes

;

Soanecennansa eas

OZ NEES POOUS"EPESERZ & P

Bonneys Camp South

POZO- TEES PNK TEP ESS ES

e Ed o<75 oo =< 7So

‘

8 Bate

| S6eW

140 p — 20

129 | Em Diptera 7

2 100 a ° +15 P

a 80 11,8

z 710 G

z 60 yo 2)

8 4 5

< | al all.

0 1 fhidtis Oe enna

FOZOT NESS EVO MESSE 4 = 4 =

oh bs 8 |

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Transactions of the Royal Society of S. Aust. (2005), 129(1), 39-42.

THE KIMBA METEORITE: AN (H4) CHONDRITE FROM SOUTH AUSTRALIA

by G. Horr!, A. PRING!? & M. ZBIK!3"

Summary

Horr, G., PRING, A. & ZBIK, M. (2005). The Kimba meteorite: An (H4) chondrite from South Australia. Trans.

R. Soc. S. Aust. 129(1), 39-42, 31 May, 2005.

The Kimba meteorite is a single stone of 1.492 kg found between 5 and 10 km south of Kimba, South Australia

(33° 13'S 136° 25’ E). It consists of olivine (Fay). n = 15), low-Ca pyroxene (Fsi67Woi2 n = 10), high-Ca

pyroxene and troilite. All of the iron-nickel metal and most of the troilite appear to have oxidized to a mixture

of iron and iron nickel oxides and hydroxides. Based on its texture and mineral chemistry, the Kimba meteorite

is classified as an H4 chondrite of shock stage S1 and weathering grade W4.

Key Worpbs: Kimba, meteorite, chondrite.

Introduction

A single mass of the Kimba meteorite was found by

Mr Lyall Cliff of Kimba more than 20 years ago, on

one of his two properties (sections 43 and 21) south

of Kimba, South Australia (33° 13’ S 136°

25’ E) (Fig 1). The meteorite lay unrecognized on the

windowsill in a shed on Mr Cliff's property until

November 1995, when Byron Smith, a member of a

local farming family, noticed the stone and suggested

that it was a meteorite. Mr L. M. Smith brought the

stone to the South Australian Museum, where the

identification was confirmed. Unfortunately, Mr Cliff

was unable to remember exactly when he picked up

the stone, or on which of his two properties it was

originally found: they are both south of the township

of Kimba and about 5 km apart.

Under legislation enacted by the Government of

South Australia, all meteorites found in the state are

the property of the Crown. The Board of the South

Australian Museum decided to reward both Mr Cliff

and Byron Smith for their part in finding this

meteorite. In accordance with the guidelines on

meteorite nomenclature, the meteorite has been

named Kimba after the geographical locality closest

to its site of discovery. The meteorite’s name and

petrological classification have been approval by the

Meteorite Nomenclature Committee.

Physical Description

The Kimba meteorite resembles in outline one half

of a rectangle broken upon a diagonal (Fig. 2). The

' Department of Mineralogy, South Australian Museum, North

Terrace, Adelaide, South Australia 5000.

? Department of Geology and Geophysics, University of Adelaide,

Adelaide, South Australia 5005,

‘Jan Wark Research Institute, University of South Australia,

Mawson Lakes, Adelaide, South Australia 5095.

* Author to whom all correspondence should be addressed

triangular section measures about 11 x 11 x 15 cm,

and is about 6 cm deep. The stone has a deep fissure

caused by weathering, with subsidiary cracks

branching from it, but the whole is very coherent and

solid. The surface is dark reddish brown and rough

because of the adherence of grains of quartz. In the

surface oxidation crust, a number of black spots or

blisters up to 2 cm across have developed, probably

caused by weathering and oxidation of iron-nickel

metal and troilite (Fig. 2).

Petrographic Features

One corner of the stone was sawn from the mass

and a polished thin section was prepared from the

off-cut. This section was used for the petrographic

examination and for electron microprobe analysis.

The interior of the meteorite is dark-grey, and

chondrules and some troilite are apparent under low

magnification. The chondrules and chondrule

fragments are partly recrystallised and have well-

defined boundaries, so are recognisable even without

using crossed polars. They are typically less than 0.5

mm in diameter but some chondrules measure up to

2 mm. Olivine and pyroxene grains up to 0.5 mm

across, along with chondrules and their fragments,

are embedded in dark, opaque, ferruginous matrix.

Troilite occurs as finely disseminated grains

throughout the matrix.

Several varieties of chondrules are apparent.

Barred olivine chondrules (BO type of Wasson

(1993)) are most distinctive in the studied thin

section. They appear as skeletal olivine with thin

lamellae of turbid, glassy mesostasis with a Na-

dominant feldspar composition. Chondrules of a

second type, radial pyroxene (RP), are also present,

and several of these display a cryptocrystalline

structure with wavy extinction. A number of granular

olivine - pyroxene (GOP) chondrules and porphyritic

pyroxene (PP) chondrules are present, containing

40 G. HORR, A. PRING & M. ZBIK

136° 137° 138° 139°

SOUTH AUSTRALIA PORT AUGUSTA

33°

PORT PIRIE

3a"

Fig. 1. Map showing the location of Kimba.

bObern

‘ \ | { j i | { | |

Fig. 2. Photograph of the Kimba meteorite, showing the deep weathering cracks and the large blisters on the surface.

THE KIMBA METEORITE: AN (H4) CHONDRITE FROM SOUTH AUSTRALIA 41

large, euhedral, polysynthetically twinned pyroxene

grains. A single porphyritic olivine (PO) chondrule

is present in the thin section and it displays coarse,

euhedral olivine grains with turbid intergranular

mesostasis. Cryptocrystalline pyroxene chondrules

diameter. In addition, there are lithic fragments up

to 4 mm in diameter of sub-rounded aggregates of

olivine crystals and microcrystalline groundmass.

The olivine grains in Kimba display normal

extinction (unstrained) and are unaltered but show

irregular cracks that are stained by iron oxide.

While some troilite is evident by reflected light

microscopy, all of the iron-nickel metal appears to

have been oxidized.

Mineralogy

Compositions of the silicate minerals were

determined using an electron microprobe

(CAMECA SX51 with Moran analysis package) at

the University of Adelaide Centre for Electron

Microscopy and Microstructure Analysis (now

Adelaide Microscopy). Analyses were made using

an accelerating voltage of 15 keV, a sample current

of 20 nA, and beam width of 0.1 Um.

Olivine in the Kimba meteorite is equilibrated,

with a mean fayalite content of Faio.2+03

(n=15){Mg/(Mg+Fe)}. The orthopyroxene shows

only small variations in chemical composition, with

a mean ferrosilite content of Fsi67::3 (n=10)

{Mg/(Mg+Fe)} and a wollastonite content of 1.2

mol% (n=10){Ca/(Cat+Mg+Fe)}. The composition

of the high-Ca pyroxene varies from grain to grain.

The glassy mesostasis has _ feldspar-like

composition (approximately Abs2Ani3Ors) and is

probably not original as the meteorite is heavily

weathered.

Classification

The Kimba meteorite has been classified as an H4

chondrite. The olivine (Fa,y..) and low-Ca pyroxene

(Fs\,7) compositions are within the range of the H

chondrites (Keil & Fredriksson 1964). The well-

defined chondrule boundaries, equilibrated olivine

and low-Ca pyroxene compositions, abundance of

high-Ca pyroxene which varies in composition, and

feldspar-like mesostasis predominantly consisting

of microcrystalline material, suggest that the Kimba

meteorite belongs to the type 4 classification of Van

Schmus & Wood (1967). The wollastonite content

in the low-Ca pyroxene lies in the higher range of

H4 chondrites (Scott et al., 1986).

Olivine and pyroxene crystals in the Kimba

chondrite display irregular fractures and sharp

optical extinction, which all indicate that the

meteorite does not seem to be shocked after

metamorphism. According to the classification

scheme of Stoffler e¢ a/. (1991), the shock facies is

estimated to be S1, i.e. unshocked. Near complete

oxidation of metal and troilite, but no alteration of

silicates in the meteorite indicates weathering state

W4 on the classification scheme of Wlotzka (1993).

Related Meteorites

Kimba is the eighth H4 chondrite to be reported

from South Australia, but the only one from the

Eyre Peninsula. Four of the others, Witchelina,

Kittakittaooloo, Coonama and Myrtle Springs, were

found in the northeastern region of South Australia

and have recently been reviewed (Zbik and Pring

2004). The other three H4 chondrites (Cook 004,

Cook 007 and Cook 009) are all from the Cook area

of the Nullarbor Plain, some 600 km to the west of

Kimba (Grady 2000, Bevan and Pring 1993). It is

unlikely that any of the other South Australian H4

chondrites would be part of the same fall as Kimba,

as the nearest site, Myrtle Springs, is some 330 km

to the North.

The Eyre Peninsula of South Australia is one of

the most productive areas of the state for meteorite

finds. This is a product of both its dry climate and

the extensive clearing of the land for cropping (see

Wallace and Pring (1991) for a brief summary).

Four other H-group chondrites have been found in

the area: Buckleboo (H6), Kaldoonera Hill (H6),

Kappakoola (H6) and Kielpa (H5). The distinction

between type 4 and 5 in the Van Schmus and Wood

classification is based on the abundance of

clinopyroxene, with type 5 chondrites having only

very minor amounts, While classification into the

petrological types can be subjective, we think that it

is very unlikely that Kimba and Kielpa are related,

even thought the average olivine compositions are

somewhat similar (Fais» for Kielpa and Fai9.2 for

Kimba).

Acknowledgments

The authors are thankful to Ben McHenry for

drafting Fig. | and Trevor Peters for the photograph

of the meteorite (Fig. 2). We also wish thank Mr

Angus Netting and Mr John Terlet, of Adelaide

Microscopy in the University of Adelaide, for

assistance with the electron microprobe analyses.

Dr W. D. Birch and Dr A. W. R. Bevan provided

constructive comments on an earlier version of this

manuscript.

42 G. HORR, A. PRING & M. ZBIK

References

BevAN, A. W. R. & PRING, A. 1993. Guidelines for the

naming of new meteorite finds from the Nullarbor

Region, South Australia. Meteoritics, 28, 600-602.

Grapy, M. M. 2000. Catalogue of meteorites. 5th edition.

Cambridge University Press London. 690pp.

KeIL, K. & FREDRIKSSON, K. 1964. The iron, magnesium

and calcium distribution in coexisting olivines and

rhombic pyroxenes of chondrites. J. Geophys. Res. 69:

3487-3515.

Scort, E. R. D., TAYLor, G. J. & KEIL, K. 1986. Accretion,

metamorphism, and brecciation of ordinary chondrites:

Evidence from petrologic studies of meteorites from

Roosevelt County, New Mexico. Proc Lunar Planet. Sci.

Conf. 17th, E115-E123.

STOFFLER, D., KEIL, K. & Scott, E. R. D. 1991. Shock

metamorphism of ordinary chondrites. Geochim.

Cosmochim. Acta. 55: 3845-3867.

VAN ScHMus, W. R. & Woop, J. A. 1967. A chemical-

petrologic classification for the chondritic meteorites.

Geochim. Cosmochim. Acta 31: 747-765.

WALLACE, M. E. & PRING, A. 1991. The Mangalo meteorite,

a new (L6) olivine-hypersthene chondrite from South

Australia. Trans. R. Soc. S. Aust. 115, 89-91.

Wasson, J. T. 1993. Constraints on chondrule origins.

Meteoritics 28: 14-28.

WLOTZKA, F. 1993. A weathering scale for the ordinary

chondrites. Meteoritics: 28, 460.

ZBIK, M. & PRING, A. 2004. The Myrtle Springs meteorite:

a chondrite (H4) from South Australia. Trans. R. Soc. S.

Aus. 128, 33-36.

Transactions of the Royal Society of S. Aust. (2005), 129(1), 43-48.

BREEDING BIOLOGY OF LITORIA MICROBELOS (COGGER)

(ANURA: HYLIDAE)

by M. Anstis* & M. J. TYLER*

Summary

Anstis, M. & Tyrer, M. J. (2005) Breeding biology of Litoria microbelos (Cogger) (Anura:Hylidae). Trans. R.

Soc. S. Aust. 129(1), 43-48, 31 May, 2005.

The embryonic and larval development of the Javelin Frog, Litoria microbelos, breeding sites and form of the

egg mass are described. At 0.78 mm diameter, the ovum is the smallest of all currently known Australian frog

species. Two clutches included 259 and 277 eggs respectively. The tadpole can be distinguished by its very small

size, body shape and features of the oral disc.

Key Worps: Litoria microbelos, hylid, clutch size, embryonic development, larval development, habitat.

Introduction

The Javelin Frog, Litoria microbelos (Cogger) is

the smallest known species of Australian hylid frog

and its distribution extends from coastal north-

western Australia to north-eastern Queensland

(Barker et al., 1995), where breeding occurs in

ephemeral water bodies during the wet season. A

brief account of the breeding behaviour and early

development is given by Tyler ef a/. (1983), but no

material beyond stage 27 (Gosner, 1960) was

available for that study and no illustrations were

provided. The present paper provides a more

complete description of the embryonic and larval

development of this species.

Materials and Methods

Embryos: Two egg clutches laid by two females

collected near Coolalinga, Northern Territory were

studied. Two females and two calling males were

collected individually at 2030 hrs on 5. ii. 2003 from

the edge of a shallow, temporary flooded depression

in grass near a larger water body. Each male was

placed with a gravid female in a separate inflated

plastic bag containing rainwater and some grasses.

Amplexus took place and egg clutches from each

pair were raised to hatching stages in shallow water

to a depth of 3 cm, at 25-27°C.

Embryos between stages 11 and early 25 were

studied using the staging descriptions of Gosner

(1960), and Anstis (2002) between stages 19-25.

Larvae: Tadpoles studied included those raised

from the above clutches and samples collected at

Marrara near Darwin and Coonjimbah Billabong

“ 26 Wideview Rd., Berowra Hts, N.S.W. 2082, Australia

' Department of Environmental Biology, University of Adelaide,

S.A. 5005, Australia.

near Jabiru, N.T. Larvae collected in the field were

maintained to metamorphosis to confirm identity.

Tadpoles were raised outdoors in available

sunlight/shade conditions within containers of pond

water to a depth of 18 cm, on a substrate of sand and

rooted vegetation. They were fed on goldfish flakes,

frozen lettuce and small protein food sticks once they

reached stage 35, Specimens were measured with an

ocular micrometer attached to a Wild M5

stereoscopic microscope and vernier calipers. Live

specimens were examined under the microscope and

then preserved in 4% phosphate-buffered formalin at

various developmental stages (Gosner, 1960).

Measurements of embryonic and larval stages are

presented in Tables | and 2. Descriptive terminology

follows Anstis (2002). Illustrations were made by

Anstis, with the aid of a drawing tube attached to the

microscope.

Morphometric measurements in mm (see Anstis ef

al., 1998) were taken on a small sample of stage 36-

39 tadpoles anaesthetised in 1% chlorbutol solution.

Abbreviations are as follows:

Lateral view: TL = total length; BL = body length;

BD = body depth; BTM = depth of tail musculature

at base of tail; TD = maximum tail depth; DF = depth

of dorsal fin (at TD); VF = depth of ventral fin (at

TD); TM = depth of tail musculature (at TD); SS =

snout to uppermost corner of opening of spiracle; SE

= tip of snout to anterior rim of eye; SN = tip of snout

to anterior rim of naris; ED = eye diameter.

Dorsal view: BW = body width; EBW = body

width at level of eyes; BTMW = width of tail

musculature at base of tail; IO = inter-orbital span

measured between inner edge of each eye; EN =

anterior edge of eye to posterior edge of naris; IN =

internarial span, measured between inner rim of each

naris.

Ventral view: ODW = maximum width of oral

disc.

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BREEDING BIOLOGY OF LITORIA MICROBELOS (COGGER) (ANURA: HYLIDAE) 45

TABLE 2. Measurements of embryos, mean with ranges in

parentheses in mm.

Stage n Ovum Capsule

11 11 0.78 (0.77-0.82) 2.84 (2.05-3.28)

12 7 0,81 (0.77-0.86) 2.65 (2.05-3.11)

17 8 1.57 (1.35-1.72)

Total Length Body Length

19 14 2.89 (2.7-3.11) 1.8 (1.72-1.88)

21 15 3.71 (3.6-3.85) 1,86 (1.8-1.96)

22 9 4.0 (3.85-4.18) 1.88 (1.8-1.92)

24 8 4,5 (4.34-4.67) 1.91 (1.84-1.96)

Fig. |. a—Stage 17, b— Stage 19, c — Stage 21.

Results

Breeding Sites

Adult males call from grass stems, low branches of

shrubs, and on the ground beside temporary pools,

flooded ditches and larger semi-permanent ponds,

swamps and billabongs. At the Coolalinga site, many

males were calling in a swampy area by 1600 on 5

February, 2003 after a heavy afternoon storm. That

night, most frogs were active near recently formed

small shallow puddles in grassland beside the main

swamp. Four calling males were on a single small

grass tussock and there were numerous such tussocks

(and low shrubs) supporting frogs around the edges

of the swamp and flooded areas nearby.

Oviposition and embryonic development

Amplexus is axillary. Oviposition occurred after

2130 during the night of 5 February but was not

observed. The egg clutches contained 277 and 259

eggs. Most eggs were laid singly on the bottom of the

bag, but some were adhering in small clusters of 2-

10 eggs, and others were attached to vegetation or

fine twigs.

Early development is quite rapid and embryos first

examined at 0930 on 6 February, were at stages 11

and 12. The ovum is very small (mean diameter 0.78

mm, stage 11) and at stage 11, the animal pole is

dark brown and the yolk is white. The capsule is a

single fluid layer, initially spherical but becoming

more fluid and expanded as the embryo develops.

Measurements of embryos are provided in Table 1.

At 1210 on 6 February, embryos were at stage 13

and appeared dark brown macroscopically but lighter

brown under the microscope (x6 magnification). By

1530 most embryos were at early stage 17: head

broad in dorsal view, arched in lateral view; tail bud

points upwards; gill arches quite prominent; adhesive

organs well developed; stomodaeal pit visible;

hatching gland lightly pigmented. By 2000 all

embryos had reached stages 17-18. Details of

embryonic development from hatching are presented

in Table 1-3 and Figure 1.

Larvae

The period of greatest growth is between stages 25

and 30, when TL and BL measurements treble. From

40. BL increments are smaller per stage (0.5 mm or

less). A composite description of fully grown live

tadpoles at stages 36-39 is given, with some

comparison to stage 25,

Body: small, cylindrical, slightly wider than deep;

snout broadly rounded in dorsal view, rounded in

lateral view; eyes lateral, prominent and with slight

dorsal tilt, iris mostly dense gold with thin copper

ring around pupil; nares closer to snout than to eyes,

widely spaced, directed dorsally, quite large and

cavernous relative to size of body; spiracle well

below body axis, opens dorsoposteriorly, posterior to

midpoint of body; vent tube dextral, short, opens

midway up ventral fin (type a, Anstis 2002).

Tail: fins shallow to moderate, dorsal fin begins at

base of body, arches acutely to greatest depth

anterior to midpoint and tapers gradually to narrowly

rounded tip; ventral fin mostly fairly shallow with a

slight arch near midpoint, then tapers evenly to tip.

Musculature moderate and tapers to a fine point.

Pigmentation in Life

Dorsal view: broad, dark brown band extends from

snout, between nares, over chondrocranium and

along vertebral region; head region less pigmented

on either side of band then darker again at each side

anterior to eye, fine melanophore flecks all over

anterior half of body; pigment less dense over

46 M. ANSTIS & M. J. TYLER

TABLE 3. Measurements of larvae and metamorphs; mean

with ranges in parentheses in mm.

Stage n Total Length Body Length

25early 17 5.05 (4.71-5.33) 2.16 (2.05-2.29)

25 late 17 8.05 (6.27-9.17) 3,54 (2.89-4.02)

30 2 15.45 (15.13, 15.77) 6.35 (6.27, 6.44)

31 5 16.38 (15.13-16.59) 6.47 (6.27-6.60)

33 2 17.30 (16.74,17.87) 7.0 (6.60,7.40)

34 1 16.58 6.92

35 7 18.34 (17.54-9.32) 7.11 (6.92-7.24)

36 6 19,27 (18.5-20.0) 7.37 (7.24-7.56)

37 4 19,90 (18.83-21.0) 7.42 (7.32-7.56)

38 2 20.75 (20.5, 21.0) 7.84 (7.64, 8.05)

39 5 22.65 (20.8-23.7) 7.98 (7 40-8.21)

40 3 21.40 (20.2-22.8) 7.72 (7.40-8.05)

41 1 22.2 7.88

44 3 10.67 (9.48-12.39) 7.18 (6.76-7.56)

45 2 7.80 (7.56, 8.05) 7.12 (7.08, 7.16)

46 11 7.46 (6.44-8.21)

TABLE 4. Measurements of morphometric characters (mm)

for one larva at Stage 36 and 39.

Character Stage 36 Stage 39

TL 20.4 23.7

BL 8.05 8.21

BD 3.7 4.02

BW 4.02 4.5

EBW 3.54 3.7

BIM 1.61 1.61

BIMW 1.61 1.61

TD 4.34 4.5

DF 1.77 2.09

T™ 1.45 1.28

VF 1.13 1.13

10 1.93 1.93

IN 0.8 0.8

EN 0.96 0.88

N 0.32 0.32

SS 5.15 4.99

SN 0.8 0.64

SE 1.93 1.77

ED 1.28 1.28

ODW 2.09 2.09

abdomen, where it becomes somewhat speckled on

either side of darker band. Dorsal surface of tail

musculature patchy black and gold, at times

appearing banded.

Lateral view: sides of body less pigmented over

upper half of abdomen (intestines partly visible) with

some gold clusters, merging with opaque

silver/white over lower half. Dark lateral stripe from

snout to eye, broadens posterior to eye over anterior

upper half of abdomen, then becomes less dense and

mostly obscured by a fine layer of golden

iridophores. Gold continues over branchial region

with clumps of melanophores and gold clusters

Fig. 2. a — Stage 37 (dorsal view), b — Stage 37 (lateral

view).

Fig. 3. Oral Disc.

beneath eye. Dorsal surface of limbs with patches of

melanophores and gold; a line of melanophores and

gold borders each toe. Sides of tail musculature have

a mostly continuous dark stripe down the middle

bordered above and below by a clearer or non-

pigmented stripe highlighted with fine gold clusters.

Fins densely speckled with small black clumps of

melanophores and fine gold clusters of iridophores,

otherwise clear.

Preserved specimens at stage 25 have only stippled

melanophores over most of the darker dorsal areas,

while larvae in later stages from about 35 have dense

dark areas, readily visible once the gold iridophores

have gone.

Ventral view: dense, opaque copper sheen over

abdomen and clear over anterior half apart from a

few fine gold clusters.

The copper sheen on the ventral surface is thin and

patchy in live individuals at stage 25 and the dark

stripe down the lateral surface of the tail musculature

is less defined.

Oral Disc (Fig. 3, specimen at stage 39): directed

ventrally, quite wide (e.g. ratio ODW/EBW = 0.59,

stage 36); wide anterior medial gap in marginal

papillae, one row of papillae on either side of gap

increasing to 2-3 laterally and one row around

BREEDING BIOLOGY OF L/TORIA MICROBELOS (COGGER) (ANURA: HYLIDAE) 47

posterior margin with a few additional submarginal

papillae in some specimens.

Two upper and three lower tooth rows; A! is the

longest row, A? with narrow but distinct medial gap.

Rows P!24 entire and P? may be very slightly shorter

than P! and P?. Jaw sheaths medium (Anstis, 2002),

upper sheath with long lateral processes, lower

sheath quite acutely V-shaped.

Metamorphosis

It was not possible to maintain the larvae hatched

from eggs until metamorphosis. Larvae collected

from the field at stages 38-40 on 6 February, 2003

began to metamorphose on 10 February and strongly

resemble the adult in shape. Dorsum yellow-gold

with bronze sheen developing, iris golden and a

small silver line across base of each finger and toe

disc; venter mostly opaque white. Lateral stripes of

adult not yet defined. Six anaesthetised metamorphs

from Marrara at stage 46 have a mean length of 7.87

mm (7.24 — 8.21) and five preserved specimens from

Coonjimbah Billabong have a mean of 6.98 mm

(6.44 — 7.72).

Larval Behaviour

L. microbelos tadpoles have been observed most

commonly on the substrate in captivity, but will at

times range through midwater regions of the water

column, In the field, they are often found on the

substrate in shallow water at the sides of ponds or

flooded ditches, hidden amongst vegetation. They

have been observed grazing on the substrate, over

rocks and on vegetation.

Discussion

Breeding Sites

Adult Litoria microbelos are abundant during the

wet season, frequently calling after rain during late

afternoon and night. Males often select very small,

shallow water-filled depressions to call from, many

of which dry up during the heat of the next day or

two. If breeding takes place in these sites, many

embryos would not survive unless adequate daily

follow-up rain occurred. However, large numbers of

calling males are also found around the edges of

larger billabongs, swamps and flooded ditches, and

populations are obviously thriving in many areas

observed from near Darwin to Jabiru. Tyler e¢ al.

(1983), state that males only call from the base of

grass tussocks if males of bicolor are absent. In the

presence of L. bicolor the L. microbelos call from

higher positions.

Oviposition and embryonic development

Although oviposition was not witnessed, it appears,

from the position of the eggs when found, that they

are laid in small groups and either become attached

to vegetation or are scattered by the adults over the

substrate. The ova measured for this study had

already reached stages 11-12 and, as a very small

increase in diameter occurs from about stage 10, it is

likely that the diameter of ova in stages 1-9 is slightly

less than that given here. However, even at stage 11,

this species has the smallest mean diameter of all

known Australian frog species, the nearest being

Crinia tinnula (0.85 mm, stage 10, Anstis, 2002).

The combination of the extremely small size of

hatchlings at stage 19 (mean TL = 2.89), short tail

and very broadly rounded tail tip, help to

distinguish this species from all other sympatric

hylid embryos at this stage (Anstis unpubl., Anstis,

2002). Embryonic development to hatching at stage

19 was reached at a minimum of 34 hours 15 min

after the eggs were laid. This is quite rapid in

comparison to that known for other sympatric

species such as Litoria rubella (stage 20, after 72

hrs at 30°C, Tyler et al., 1983), Uperoleia inundata

(stage 20, after 48 hrs at 27-28°C, Anstis, unpubl.)

and Crinia bilingua (stage 20, after 48 hrs at 25-

27°C, Anstis, unpubl.). However, embryos of the

sympatric species, Limnodynastes ornatus, hatched

at stage 20 after only 18 hours at 30°C (Tyler ef al.,

1983). A mean of 4.1 mm (4.0-4.2) for embryos at

stage 24 is given by Tyler e¢ al. (1983), similar to

the mean of 4.5 for specimens in the present study

(Table 2).

Larvae

Tadpoles raised from eggs remained in stage 25 for

at least 20 days. More observations on specimens

raised in warmer, shallower water (and other varied

environmental conditions) need to be made to test

whether this is a typical rate of development for this

species or whether development slowed as a result of

other factors during culture. Tyler ef al, (1983)

provide a mean TL of 11.4 mm for larvae at stage 27,

which is 3.35 mm longer than the mean TL of 8.05

for late stage 25 specimens in the present study.

This species can be distinguished from all other

fully grown conspecific larvae by a combination of

its very small maximum size, cylindrical body

shape, ventral oral disc, tooth row formula of 2(1)/3,

narrowly rounded tail tip (rather than finely

pointed) and pigmentation patterns. In relation to

the body types of Anstis (2002), it most closely

affiliates with a small version of type 2, (Litoria

latopalmata complex) in shape and behaviour, but

has a slightly more ventrally directed oral disc and

larger nares. Small Type 2 hylid larvae at early stage

25 commonly sympatric with L. microbelos, such as

L. nasuta, L. inermis and L. tornieri can be

distinguished from the former by their lack of lateral

stripes and more finely pointed tail tip, and

48 M. ANSTIS & M. J. TYLER

L. tornieri is easily separated from all species at stage

25, by their red tail fins (Anstis, unpublished). Type 1

sympatric hylid tadpoles also at stage 25, such as L.

rothi, L. bicolor and L. caerulea, while somewhat

similar in body shape to larvae of L. microbelos, can

be distinguished by their finely pointed tail tips,

surface-dwelling behaviour and anteroventral oral

disc, with all but L. bicolor having an LTRF (Alltig,

1970) of 2(1)/3(1). Other conspecific bottom-

dwelling larval species closer to L. microbelos in

maximum size range (prior to about stage 27) are

Crinia bilingua (Tyler et al., 1983), Uperoleia

mimula (Richards and Alford, 1993) and U.

lithomoda (Davies et al., 1986), but all of these

species have a more oval to rounded body shape,

different tail shape, a different oral disc possessing a

medial posterior gap in the marginal papillae and lack

the distinctive lateral stripes of L. microbelos larvae.

Metamorphosis

Larvae collected in the field at stages 2 30

advanced steadily to metamorphosis, especially

those at stages 38 — 40, which began meta-

morphosing four days after capture. Larval life span

could not be determined for any one group.

Acknowledgements

Research was undertaken by Marion Anstis under

the NT Frogwatch Licence no. 16881. The WWF

Frogs! Program and Stan Orchard, the co-ordinator

are gratefully acknowledged for their financial

assistance and support during this study. Jeanne

Young and the Technical Staff of the Zoology

Department at Charles Darwin University are

gratefully acknowledged for assistance with field

work, localities and materials.

References

Attia, R. A. (1970) A key to the tadpoles of the continental

United States and Canada. Herpetologica 26: 180-207.

Anstis, M. (2002) Tadpoles of South-eastern Australia: a

guide with keys. Reed New Holland.

, ALForp, R. A. & GILLEsPIE, G. R. (1998)

Breeding biology of Litoria booroolongensis (Moore,

1961), and Litoria lesueuri (Dumeril & Bibron, 1841)

(Anura: Hylidae) and comments on population declines

of L. booroolongensis. Trans. R. Soc. S. Aust. 122(1), 33-

43.

BARKER, J., GRIGG, G. C. & TyLer, M. J. (1995) A Field

Guide to Australian Frogs. Surrey Beatty, Chipping

Norton, N.S.W.

Davies, M., MCDONALD, K. R. & CorBEN, C. (1986) The

genus Uperoleia in Queensland, Australia. Proc. R. Soc.

Vict. 98(4), 147-188.

Gosner, K. L. (1960) A simplified table for staging anuran

embryos and larvae with notes on identification.

Herpetologica 16, 183-190.

RICHARDS, S. J. & ALFORD, R. A. (1993) The tadpoles of two

Queensland Frogs (Anura: Hylidae, Myobatrachidae).

Mem. Old Mus. 33(1), 337-340.

TyLer, M. J., Crook, G. A. & Davies, M. (1983)

Reproductive biology of the frogs of the Magela Creek

system, Northern Territory. Rec. S. Aust. Mus. 18(18):

415-440.

Transactions of the Royal Society of S. Aust. (2005), 129(1), 49-52.

HEPATOZOON TACHYGLOSSI SP. NOV. (HAEMOGREGARINIDAE),

A PROTOZOAN PARASITE FROM THE BLOOD OF A SHORT-BEAKED ECHIDNA,

TACHYGLOSSUS ACULEATUS

by P. CLarkK!, P. Hotz? & D. M. Spratt?

Summary

CLARK, T., HOLz, P. & Spratt, D. M. (2005) Hepatozoon tachyglossi sp. nov. (Haemogregarinidae), a protozoan

parasite from the blood of a short-beaked echidna, Tachyglossus aculeatus. Trans. R. Soc. S. Aust. 129(1),

49-52, 31 May, 2005.

Hepatozoon tachyglossi sp. nov. is described from monocytes in the peripheral blood of a debilitated short-

beaked echidna from the Healesville region of Victoria. Of the Hepatozoon Miller, 1908 species known to occur

in Australian native mammals, all of those in marsupials occur in erythrocytes and only H. muris from

introduced and native rodents oceurs in monocytes. H. tachyglossi is distinguished from H. muris by its larger

size and the lack of a capsule.

Key Worps: Hepatozoon tachyglossi, new species, echidna, Tachyglossus aculeatus.

Introduction

The blood of the short-beaked echidna,

Tachyglossus aculeatus (Shaw & Nodder, 1792), has

yielded relatively few haemoparasites. Those

identified have been restricted to the Piroplasmidae

and include Babesia tachyglossi (Backhouse &

Bolliger, 1959) and Theileria tachyglossi (Priestly,

1915) (Backhouse & Bolliger 1957, 1959;

Mackerras 1959). We report a novel species of

Hepatozoon from monocytes in the peripheral blood

of a short-beaked echidna from the Healesville

region of Victoria and compare the species with other

members of the genus previously described from

Australian native mammals.

Clinical history and methods

A male, juvenile, short-beaked echidna was

presented to the veterinary service of Healesville

Sanctuary. The animal had several injuries including

a damaged tongue and a fractured humerus. During

captive management and treatment, the animal

developed severe dyspnoea and a mucopurulent

nasal discharge. It was euthanased after a poor

response to treatment with antimicrobial drugs and

supportive therapy.

A sample of blood was collected ante-mortem from

the bill sinus. The morphology of haematological cells

in a blood film, stained with Wright’s and Giemsa

stains, was assessed by light microscopy. Tissue

School of Clinical Sciences, Division of Veterinary and

Biomedical Sciences, Murdoch University, South St, Murdoch,

Western Australia, 6150.

? Healesville Sanctuary, P.O. Box 248, Healesville, VIC, 3777.

‘CSIRO Sustainable Ecosystems, GPO Box 284, Canberra,

ACT, 2601

samples of bone marrow, liver and spleen, were

collected post-mortem and fixed in 10 percent

buffered formalin. The tissues were processed using

standard histological methods; sections were stained

with haematoxylin and eosin stains and examined

using light microscopy. The organisms and their host

cells were digitally photographed. A stage micrometer

was photographed at the same magnification and used

to insert scale bars to all micrographs. The micrometer

was also used for all measurements which are

presented in microns as the mean + standard deviation

followed by the range in parentheses.

Results

Intracellular organisms were observed within

leukocytes in the peripheral blood (Fig. 1). These

were observed in 24/50 monocytes but were not

observed in any granulocytes.

Hepatozoon tachyglossi sp. nov.

(FIG. 1)

Host

Tachyglossus aculeatus.

Location

Healesville, Victoria.

Type

Slide in South Australian Museum No. 28751.

Typically the organisms were oval to elongate in

shape but were quite pleomorphic with some

pyriform and round forms observed. The organisms

were 9.9 + 1.4 (7.8-12.4) in length and 4.7 + 0.7 (3.8-

5.9) in width (n = 18). Most organisms had an

eccentric, subterminal nucleus and some exhibited a

50 P. CLARK, P. HOLZ & D. M. SPRATT

Fig. 1. Examples of Hepatozoon tachyglossi sp. nov. within monocytes from the peripheral blood of a short-beaked echidna.

Bar = 10 um.

small amount of punctate, dark brown — black

pigment in the cytoplasm. No distinct capsule was

observed and there was only a subtle difference in

colour between the cytoplasm of the organism and

the cytoplasm of the cell. Only one organism was

evident per cell; in some cases this caused

displacement of the host cell nucleus. No consistent

changes in cell morphology were evident. No

extracellular organisms were noted. Examination of

histological sections of bone marrow, liver and

spleen did not reveal schizonts.

Discussion

The organism described is morphologically similar

to a species of Hepatozoon Miller, 1908, the only

coccidian genus that inhabits the blood of mammals.

Desser (1990) highlighted the taxonomic confusion

arising from the problem of differentiating species of

Hepatozoon from that of Haemogregarina

Danilewsky, 1885 on the basis of gamonts in the

blood of the vertebrate host. Differentiation was

based primarily on the size of the oocysts and the

presence or absence of sporocysts in the invertebrate

definitive host. Desser (1990) tested and confirmed

the hypothesis of Landau ef al. (1972) that tiny cystic

stages (cystozoites) in the liver and lungs represent a

common feature in all species of Hepatozoon and, as

a consequence, differentiation of the genera

Hepatozoon and Haemogregarina based solely on

the stages in the vertebrate host is possible. These

findings further support the evidence of Landau ef

al. (1972) that transmission of species of

Hepatozoon may be by predation, as well as by

ingestion of the infected arthropod vectors.

The genus has not been reported previously in

HEPATOZOON TACHYGLOSSI, SP. NOV. IN AN ECHIDNA 51

Fig. 2. Examples of Hepatozoon muris from the blood of

Rattus lutreolus. Bar = 10 um.

monotremes. However, H. peramelis (Welsh &

Dalyell, 1909), H. dasyuroides Mackerras, 1959, H.

dasyuri (Welsh, Dalyell & Burfitt, 1909), H. petauri

(Welsh & Barling, 1909) and H. pseudocheiri

Mackerras, 1959, have been described from

Australian marsupials (see Mackerras, 1959). In

addition, O'Donoghue and Adlard (2000) reported

several unidentified species of Hepatozoon from

Australian marsupials. All reports have recorded the

parasite within erythrocytes or ‘free’ in the blood

(Mackerras 1959; Speare et al. 1984; Bettiol e¢ al.

1996). Also, H. muris (Balfour, 1906) has been

recognised in both the introduced Rattus norvegicus

(Berkenhout, 1769) and R. rattus (Linneaus, 1758),

and the native rodents, R. fuscipes (Waterhouse,

1839) and R. sordidus (Gould, 1858) (Mackerras

1959; O’Donoghue & Adlard 2000). Hepatozoon

muris, in contrast to the species reported in

marsupials, infects leukocytes (Soulsby 1982).

The organisms in the current study were oval to

elongate in shape, as are many species of

Hepatozoon, and were similar in size to H. peramelis

(9.0 — 10.0 um by 3.0 — 3.5 um; Welsh & Dalyell

1909) and slightly larger than H. petauri (7.5 —

8.0 um by 3.5 — 4.0 lm; Mackerras 1959), H.

pseudocheiri (8.0 — 13.0 um by 1.5-3.0 um;

Mackerras 1959), and Hepatozoon sp (8.7 + 0.2 um

by 2.1 + 0.4 um, Bettiol et al. 1996). In contrast,

H. dasyuroides (12 — 13 tum by | — 2 um; Mackerras

1959) and H. dasyuri (12 um by 4 [tm; Welsh Dalyell

& Burfitt 1909, 1910) are longer, narrower parasites.

Hepatozoon tachyglossi sp. noy., like H. muris,

occurs in the monocytes of its host. However,

H. muris is a smaller (7.0 — 8.0 [tm by 3.0 — 3.5 um;

Mackerras, 1959) more ovoid, and typically less

morphologically variable parasite, than H.

tachyglossi (Fig. 2). Additionally, H. muris has an

eosinophilic capsule that was not evident in

A. tachyglossi.

Typically, the definitive hosts and vectors of

species of Hepatozoon are blood feeding arthropods.

The mite Laelaps echidninus Berlese, 1887 fills this

role for H. muris. In the current case, ticks identified

as Aponomma concolor Neumann, 1899, were

evident on the echidna but were not examined for

sporocysts and could not be proven to be the

definitive host.

Although the life history of this species of

Hepatozoon remains unknown, we consider that

specific status is warranted on the basis of

morphological features and the occurrence of the

parasite in monocytes of the monotreme,

Tachyglossus aculeatus.

The effect on the host was not determined due to

the other pathological processes and the animal’s

debilitated state may have allowed the organism to

proliferate. Further work to identify the prevalence

of this organism in short-beaked echidnas, assess

any pathogenic effects on the animal and

phylogenetic studies to determine its relationship to

other species of Hepatozoon in Australia need to be

undertaken.

Acknowledgements

We thank Dr Peter O’Donoghue for sharing his

knowledge of protozoan organisms with us and two

anonymous referees whose comments improved the

presentation of the manuscript.

References

Backnouse, T. C., & BOLLIGER, A. (1957) A piroplasm of

the echidna (7achyglossus aculeatus). Aust. J. Sci. 19,

24-25.

& (1959) Babesia tachyglossi n. sp.

from the echidna Zachyglossus aculeatus. J. Protozool. 6,

320-322.

BeETTIOL, S. S., GoLDsmID, J. M., Le, D. D. & Driessen, M.

(1996) The first record of a member of the genus

Hepatozoon in the eastern barred bandicoot (Perameles

gunnit) in Tasmania. J. Parasitol. 82, 829-830.

Desser, S. S. (1990) Tissue “cysts” of Hepatozoon

griseisciuri in the grey squirrel, Sciurus carolinensis: the

significance of these cysts in species of Hepatozoon. J.

Parasitol. 76, 257-259.

Lanbau, I., MICHEL, J. C. & CHasaup, A. G, (1972) Cycle

biologique d’Hepatozoon domerguei: discussion sur les

caractéres fondamentaux d’un cycle de Coccidie. Z.

Parasitenk, 38, 250-270.

MacKerras, M. J. (1959) The Haematozoa of Australian

mammals. Aust. J. Zool. 7, 105-135.

52 P. CLARK, P. HOLZ & D. M. SPRATT

O’ DONOGHUE, P. J. & ADLARD, R. D. (2000) Catalogue of

protozoan parasites recorded in Australia. Mem. Qld.

Mus. 45, 1-163.

Soutssy, E. J. L. (1982) “Helminths, arthropods and

protozoa of domesticated animals”. 7th ed. (Bailliére

Tindall, London.)

SPEARE, R., HAFFENDEN, A. T., DANIELS, P. W., THOMAS, A.

D., & SEAWRIGHT, C. D. (1984) Diseases of the Herbert

River ringtail, Pseudocheirus herbertensis, and other

North Queensland rainforest possums. pp. 283-302 In

Smith, A. P. & Hume, I. D. (Eds.) “Possums and

Gliders”. (Surrey Beatty & Sons, Chipping Norton.)

WELSH, D. A., & DALYELL, E. J. (1909) Haemogregarina

peramelis: a free haemogregarine of an Australian

bandicoot. J Path. Bact. 14, 547-549.

& Burritt, M. B. (1909) Haemogregarina

dasyuri. A preliminary note on an undescribed

haemogregarine of the Australian native cat. Aust. Med.

Congr. for 1908. 2, 333-337

(1910) Haemogregarina dasyuri: a haemo-

gregarine of the Australian native cat. J. Path. Bact. 14,

542-546.

Transactions of the Royal Society of S. Aust. (2005), 129(1), 53-58.

RECORDS OF HUMPBACK WHALES MEGAPTERA NOVAEANGLIAE

IN SOUTH AUSTRALIA

by C. M. KEMPER

Summary

Kemper, C. M. (2005), Records of humpback whales Megaptera novaeangliae in South Australia. Trans. R. Soc.

S. Aust. 129(1), 53-58, 31 May, 2005.

Opportunistic sightings and museum specimens of humpback whales (Megaptera novaeangliae) in South

Australia (n = 116) were collated to the year 2003. Records were made in all months of the year, with 57% during

June and July, Timing of this peak and the presence of neonates presumably related to the northward migration

from high latitudes to breeding grounds in the tropics. Sightings and beach-washed carcasses were widely

distributed, from Head of Bight to the Victorian border, with apparent concentrations in eastern Gulf St Vincent,

Kangaroo Island and Victor Harbor regions, and to a lesser extent in the south-east of the State and southern

Eyre Peninsula. Most sightings involved single whales or groups of two (range 1 — 4). All five carcasses were

neonates or juveniles with total lengths of 3.88 m to approx. 10 m. Estimated lengths (5 — 15 m) were available

for 17 live whales. The geological age of one museum specimen is uncertain but may predate European

settlement. This and whaling records from Fowler Bay during 1840 indicate that humpback whales have been

present, and may not have been uncommon, in South Australia since at least the early 19th century. Research is

needed to determine the relationship of South Australian humpback whales to populations migrating off the

western and eastern coasts of Australia.

Key Worps: Humpback whale, South Australia, sightings, strandings, distribution.

Introduction

Humpback whales (Megaptera novaeangliae)

occur in all major ocean basins and from the polar

regions to the tropics (Clapham 2002). Populations

in the southern hemisphere are broadly delimited by

longitudinal degrees: for the Australasian region

these are Group IV between 70° and 130° E, and

Group V between 130° and 170° E (Mackintosh

1965). These populations migrate along the west

coast of Australia, and the east coast of Australia and

throughout New Zealand, respectively (Dawbin

1966). They have different genetic make-up (Baker

et al. 1994), songs (Cato 1991; Dawbin & Eyre

1991) and migration routes (Dawbin 1966). At 1999,

the size of the Group IV population was estimated to

be 8207-13 640, increasing at a rate of about 10%

per annum (Bannister & Hedley 2001) and that of

Group V, 36004440 increasing at about 11% per

annum (Paterson e¢ al, 2001).

Humpback whales in the Australasian region were

taken by pelagic and shore-based whalers during the

19th and 20th centuries (see overviews by Bannister

& Hedley 2001 and Paterson 2001). Between the

early 1900s and 1963, most humpbacks off Australia

were taken from shore-based whaling stations along

the east and west coasts of the continent. There were

also significant catches by pelagic whalers during

the mid to late 1930s (Chittleborough 1965) and

illegal whaling by the Soviet fleet took many animals

South Australian Museum, North Terrace, Adelaide, South Australia

5000.

south of the continent until 1973 (Tormosov 1995;

Mikhalev 2000). There was no targeted, shore-based

whaling of humpback whales in South Australia

during the 20th century but some animals were taken

there opportunistically during 19th century whaling

for southern right whales (Eubalaena australis).

There are few published records of humpback

whales in South Australia and as a result, it is

generally thought that these represent vagrants.

However, in 1840 the whale ship Amazon took eight

humpback whales at Fowler Bay (Bannister 1986).

Chittleborough (1965) published two sightings — one

whale at the head of the Great Australian Bight in

August 1952 and a female with newborn calf in Gulf

St Vincent in winter 1961. Museum specimens and

beach-washed carcasses from South Australia were

reported and /or illustrated in Aitken (1971), Kemper

& Ling (1991) and Judd et al. (1992).

This study documents the geographic and temporal

distribution of sightings and carcasses of humpback

whales in South Australia using opportunistic

records from various sources. Some information is

also provided on relative age/size of animals, as well

as number of animals seen in each group. Avenues

for further research are explored.

Material and Methods

Records of opportunistic sightings (n = 109) of

humpback whales were obtained from the whale

sightings database at the South Australian Museum.

The Museum database has been compiled from

various sources, including the South Australian

Whale Centre (Victor Harbor), marine mammal

researchers, public, fishers, and the published

records listed above. Sighting records dated from

1948 to 2003. Records of carcasses and South

Australian Museum specimens (n = 7) dated from

‘before 1913’ to 2000.

Records were coded for reliability of

identification: 1 (n = 11) = photographic evidence; 2

(n = 50) = certain/probable (experienced observer or

distinctive features noted; i.e. white underside of

flukes, trailing edge of flukes scalloped, very long

flippers, dark lumps on head, small irregular dorsal

fin); 3 (n = 29) = some doubt (inexperienced

observer, not enough features recorded to be

confident of identification); 4 (n = 19) = no

description of features provided, observer

experience unknown but identification may be

correct.

Latitudes and longitudes, including the accuracy

of localities, were calculated by Museum staff and

trained volunteers. Distance from shore, number of

whales sighted and other details were sometimes

supplied by observers.

Carcasses provided accurate body length data as

well as the degree of physical maturity of the

prepared skeleton. Whale length was estimated quite

accurately for three live animals: one by the author

from photographs of the flukes and two by fishers

C. M. KEMPER

who related the length of the whale to that of the

boat. Relative size and/or rough estimates of whale

length were noted by some observers.

Results

Of the seven records of humpback whale

mortalities, two were bones of unknown age (of

which one was possibly subfossil) and therefore

no details were available of the carcasses,

circumstances or when the animals died (Table 1).

One juvenile (S12) probably floated in dead, and one

neonate (M21309) was very fresh when first

discovered on the beach (Fig. 1) and therefore was

probably alive nearby before it stranded or washed

up. M15187, a juvenile, was reported as being alive

in the vicinity for several days before washing up

dead. Two animals (M12778 and M16971) were in an

advanced state of decomposition when discovered

but because of the remote locations of the carcasses,

they could have been on the beach some time before

being discovered. Neither had characteristics of

animals that had been floating dead for some time

before washing up.

Humpback whales have been seen alive in many

parts of coastal South Australia, as well as up to 250

km offshore in the Great Australian Bight and up to

100 km offshore in the south-east of the State (Fig. 2).

TABLE |. Specimen records of Megaptera novaeangliae in South Australia. Date is when first seen or reported on beach or

in case of unknown age of material, when registered into collection.

No. Date Locality Length Sex Relative Decomposition Material at SAM

(m) Age

M149 __ before Largs Bay, Adelaide — — a unknown right and left bullae

1913 34° 49’ S, 138° 29’ E (provenance unknown)

M5120 _ before ‘west coast of SA’ a — —o unknown radius, ulna, humerus,

1944 scapula (provenance

unknown)

S12 8 Aug. 14 km E Vivonne, ~10 U juvenile — advanced, none

1984 Kangaroo Island much skin

36° 00' 00" S, 137° 19’ 30” E sloughed off

M12778 24 Oct. 12kmSW 5.53 —, neonate — quite almost full skeleton

1985 Streaky Bay, decomposed

Great Australian

Bight

32° 52’ 10” S, 134° 07' 00" E

MI15187_ 1 Mar. 3 km N Point Yorke, 7.25 male — juvenile intact but almost full skeleton,

1989 Investigator Strait starting to baleen, barnacles

35° 12’ S, 137° 12' E decompose

M16971 late Dec. 12.5 km SSE Ceduna, 7.10 as juvenile — very skull collected,

199] Great Australian Bight decomposed barnacles

32° 17'S, 133° 39’ E

M21309 27 July 7.4 km NW Elliston, 3.88 female neonate very fresh full skeleton, baleen,

2000 Great Australian Bight organs, genetics tissues,

33° 36’ 00” S, 134° 50 20" E

toxicology tissues

HUMPBACK WHALES IN SOUTH AUSTRALIA 55

Many sightings were made from land—some being

seen as close as 50 m from shore. Concentrations in

sightings are apparent for eastern Gulf St Vincent,

Kangaroo Island and Victor Harbor regions, and to a

lesser extent off the lower Eyre Peninsula and in the

south-east of the State. Humpback whales have been

seen well within the gulfs, including a reliable

record north-east of Whyalla and several north of

Adelaide. When whale movements were ‘tracked’ by

successive observations south of Adelaide, the

pattern was northbound. Local residents reported

Fig. 1. Female Megaptera novaeangliae neonate (M21309) that humpback whales were often sighted off

washed up at Anxious Bay, 7.4 km NW Elliston, SA on 27. _ Elliston, generally moving north (Marie Clark, pers.

July 2000. Photo: C. Kemper/South Australian Museum. comm.).

| |

| Z ;

South Australia |

| |

! |

| gg Head of Bight |

a — |

ms Fowder Bay |

saa: ata |

. 7 an, |

aa Wrserg iN |

he i 2 |

E P I :

vA “ee yre Peninsula aS %,

es. Cag Elliston £7 as |

~ r Pon ¢

4 Mf gt ios era

Southern Ocean a oF |

~ | emia |

« ? t

a |

. ena Th |

< '

N a

vhs Les |

3 -

0 70 140 280

ae Nautical Miles ae y A

Fig. 2. Geographic distribution of Megaptera novaeangliae in South Australia. Closed triangles = sightings reliability | and

2, closed circles = sightings reliability 3 and 4, open squares = South Australian Museum specimens, * = towns/cities

mentioned in text.

Ww ow

oun

Mild 3 and4

Old tand 2

No. records

J oF MAM J J

Month

A S ON D

Fig. 3. Month of observation for sightings and carcasses of

Megaptera novaeangliae in South Australia. Id =

reliability of identification (see Material and Methods).

*= one carcass record,

Fig. 4. Megaptera novaeangliae trapped in tuna feedlot off

Boston Island, Spencer Gulf (34° 44" S, 135° 55’ E) on 14

June 1993. Photo: Daryl Lawrence/NPWSA.

Only five of the sightings and no intact carcasses

were recorded before 1980, Between 1980 and 1989,

there were 17 sightings and three carcasses. The

remaining records (79%, n = 87) dated from 1990 to

2003. Sightings of live animals were made during all

months of the year and carcasses were recorded in

March, July, August, October and December (Fig. 3,

Table 1). Sixty (57%) of the sightings were made

during June and July. There did not appear to be a

relationship between region and month sighted—

both eastern and western parts of the State had

humpback sightings in all seasons.

Accurate data on body length and relative age

came from carcasses collected for the South

Australian Museum (Table 1). Of the five measured

specimens, two were neonatal in length (5.53,

3.88 m) and three were juveniles (7.10, 7.25, ~10

m). Estimates of body length of 17 live animals

(5-15 m) were available, mostly from boat-based

observers. (Since boating enthusiasts are usually

familiar with the length of their vessel and can

compare a whale with this, these estimates are

C. M. KEMPER

considered reasonably accurate.) The length of a live

whale entrapped in a tuna feedlot near Port Lincoln

was estimated (10 m) by the author by measuring

fluke width from photographs submitted (Fig. 4). A

number of observers reported ‘adult and calf’,

‘subadult’ or ‘large’ and ‘medium’-sized animals

travelling together but with no supporting estimation

of body length or how the relative size was

determined.

Humpback whales have been observed interacting

with southern right whales at Head of Bight during

winter, one such interaction lasting for >75 min (S,

Burnell and R. Pirzl, pers. comm.). Humpback

whales were seen during five out of seven years of

southern right whale studies at Head of Bight.

The number of humpback whales observed during

sighting events in South Australia often

accompanied reports (n = 98). The results are

summarised as follows: | animal = 44 times, 2 = 44,

=8,4=2.

Discussion

The number of records of humpback whales for

South Australia is ample evidence that this species is

more than a vagrant to the south-central coast of the

continent. In addition, sightings are made from year

to year in some places, e.g. Head of Bight. The

documenting of 19th century and recent records is

evidence that humpback whales have been present

off South Australia for more than 160 years. The

American whale ship, Amazon, took eight

humpback whales and 33 southern right whales at

Fowler Bay during an 80-day period in the winter of

1840 (Bannister 1986), An excavation of the site in

1994 identified many bones of southern right whales

but no humpback whales, although not all material

was identifiable (Kemper & Samson 1999),

Townsend (1935) compiled logbook records of

American whale ships from the 19th century and

noted only two humpback records for the waters off

southern Australia, both near Albany, Western

Australia in July. With Australian populations of

humpback whales increasing since at least the early

1980s (Bannister & Hedley 2001; Paterson er al.

2001), it is expected that sightings off South

Australia will become more frequent.

The coast of South Australia spans the longitudes

129° to 141° E, which falls within the limits of the

Group V_ population. However, the affinity of

humpback whales in that State to Group IV and/or V

populations is not known. Between the 1930s and

1960s research on great whale movements was

based largely on Discovery marking of whales in the

Antarctic and Oceania, and off Australia and New

Zealand (Dawbin 1966). This technique of studying

the movements of whales involved firing numbered

HUMPBACK WHALES IN SOUTH AUSTRALIA 57

markers into live whales then these markers were

often recovered during whaling operations. Since

there were no 20th century whaling stations along

the coast of South Australia (Findlay 2001), no

marks were recovered there. If members of both

humpback whale populations occur off South

Australia they may remain discrete (possibly Group

IV in the Great Australian Bight and Group V off the

south-east of the State) or there may be no

geographic separation. Dawbin (1966) reported

limited mixing of these populations in their

Antarctic feeding grounds.

Humpback whales occasionally calve south of

28° S on the east coast of Australia (Janetski &

Paterson 2001) but there are no known current

calving grounds off southern Australia (Bannister ef

al, 1996). Three ‘cows and calves’ were taken by the

Amazon during June to September 1840. Since no

body lengths were recorded, the ‘calves’ may have

been either neonates (possibly born prematurely on

the northward migration) or young of the previous

year. Humpback whales breed in tropical waters of

at least 25°C (Dawbin 1966) and the waters off

South Australia are temperate (i.e. 10—20°C). It is

theretore unlikely that this is an undescribed calving

ground.

In the Australasian region humpback whale

migration is segregated by age, gender and

reproductive status (Chittleborough 1965; Dawbin

1966; Bannister ef a/. 1996). On the northward

migration (mostly June to August but some as late as

October (Paterson ef al. 2001)) females with

yearling calves travel first, followed by adult males

and non-pregnant females, then lastly females in late

pregnancy. On the southward migration, which

occurs from August/September to November/

December (Paterson ef a/. 2001) females in early

pregnancy travel first, followed by immature

animals, then resting females with mature males and

lastly cows with young calves (Dawbin 1966). There

is evidence that not all females migrate north to the

breeding grounds each year (Brown ef al. 1995).

Thus it could be hypothesised that some of the South

Australian sightings are these females.

Chittleborough (1965) noted that some Group IV

humpback whales meet the south coast of Western

Australia on their northward migration, then travel

west/south-west to continue north along the west

coast of the continent. This observation, and the fact

that the majority of South Australian sightings were

in June and July, suggests that the latter are part of

the northward migration. The presence of dead

neonates in July and October is presumably linked to

the last of the northbound migration, i.e. late-

pregnancy females. Humpback whales are weaned

at lengths of 7.5 to 9 m when less than 12 months

old (Bannister et al. 1996). Of the three dead

juveniles recorded in South Australia, one was 7.2 m

(March) and the other 7.1 m (December). The March

juvenile may have been migrating north with its

mother. The timing and size of the December

juvenile suggests a southbound migration as either a

dependent calf or weanling. The August carcass of

an animal estimated at 10 m body length was

probably weaned and may have been on a northward

migration.

A small proportion of humpback whales feed

opportunistically off Australia on their migrations

north and south. Chittleborough (1965) found only

five out of 197 humpbacks sampled off the Western

Australian coast had food (small quantities) in their

stomachs. The species has been observed feeding on

the euphaustid crustacean, Nyctiphanes australis,

off eastern Tasmania and on an unknown prey off

southern New South Wales (Gill ef al. 1998). Pygmy

blue whales have been seen feeding on N. australis

during summer and autumn in the Bonney upwelling

off western Victoria and south-eastern South

Australia but as yet no humpback whales have been

seen there (Gill 2002). A humpback whale was

observed on 16 July 1993 tn Discovery Bay,

Victoria, very close to the South Australian border

during surveys for southern right whales (Kemper,

unpublished data) but this is not when upwelling

events occur.

Future research on humpback whales in South

Australia should include |) determining affinities to

existing Australian populations by genetic studies

and photo-identification (some tissues and

photographs presently available at the South

Australian Museum), 2) estimating relative

abundance and mapping offshore distribution using

platform of opportunity surveys and 3) determining

sex and relative age structure through close study

and biopsy of live animals.

Acknowledgements

Research involving opportunistic sightings of

whales relies on researchers, members of the public

and government officers to submit records. I thank

all those who have contributed, especially Stephen

Burnell and Rebecca Pirzl, Trevor Whibley, officers

of National Parks and Wildlife South Australia and

the South Australian Whale Centre. Sightings

accumulated by John Ling and Pin Needham formed

the basis of the whale records and BHP Ltd.

financed their databasing at the South Australian

Museum. Many people are thanked for their work on

that database — Jenny Mole, Helen Owens, Tanya

Taylor, Claire Taylor, Michaela Ciaglia and Tami

Stone. | am grateful to the following people for

assistance with collecting humpback whale

carcasses: Lynette Queale, Catherine Bell, Ron

58 C. M. KEMPER

Waterhouse, Ross Allen, Don Mount, Tom

Gerschwitz, David Farlam, Trevor Whibley.

Museum staff members (Jim McNamara, Terry Sim,

Lynette Queale, Peter Cockerham, Bob Hamilton-

Bruce) have played an important role in preparing

and managing the cetacean collection. John

Bannister provided helpful comments on a draft

manuscript.

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& HEDLEY, S. L. (2001) Southern hemisphere

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Transactions of the Roval Society of S. Aust. (2005), 129(1), 59-64.

A REDESRIPTION OF ODILIA EMANUELAE (NEMATODA: TRICHOSTRONGYLINA:

HELIGMONELLIDAE) FROM AUSTRALIAN RODENTS WITH A KEY AND

COMMENTS ON THE GENUS ODILIA

by L. R. SMALES*

Summary

Smaces, L. R. (2005) A redescription of Odilia emanuelae (Nematoda: Trichostrongylina: Heligmonellidae)

from Australian rodents with a key and comments on the genus Odilia. Trans. R. Soc. S. Aust. 129(1), 59-64, 31

May, 2005.

Odilia emanuelae (Mawson, 1961) is redescribed from a new host, Raffus leucopus from Queensland Australia.

A key to the genus Odilia, based on the number and size of the ridges of the synlophe is given. The relationships

of species of the genus Odilia with their murid hosts, coevolution or host switching, are discussed and

speculation as to the significance of their known biogeographic distribution is put forward.

Key Worps: Nematoda, Heligmonellidae, murid, Ra/tus, Australia, phylogeny, biogeography.

Introduction

During a study of the helminth parasites of the

Cape York rat, Rattus leucopus (Gray), specimens of

three species of trichostrongyloid nematodes,

Heligmonellidae, were encountered in the duodenum

of several hosts. They included Nippostrongylus

brasiliensis (Travassos, 1914), a cosmopolitan

species and NV. magnus (Mawson, 1961), endemic to

Australian rodents. Previously noted from Rattus

fuscipes (Waterhouse), R. sordidus (Gould) and

Melomys cervinipes (Gould) (see Smales, 1997), WN.

magnus was redescribed by Beveridge & Durette-

Desset (1992) from R. fuscipes and experimentally

infected Rattus norvegicus (Berkenhout). The third

species found was Odilia emanuelae (Mawson,

1961). Odilia emanuelae was described originally

from R. sordidus as R. conatus and R. fuscipes as R.

assimilis by Mawson (1961) but she did not describe

some features including the cuticular ridges of the

synlophe. Subsequently a brief description of the

synlophe was given by Durette-Desset (1969) on the

basis of one specimen. The additional material from

a new host allows a more detailed description to be

prepared and a key to known species of the genus

given.

The genus QOdilia has been reported from 11

endemic species of murids from Australia: in the

subfamilies Hydromyinae, Mastacomys fuscus

Thomas, Melomys burtoni (Ramsay), Melomys

cervinipes, Mesembryomys gouldii (Gray),

Pseudomys higginsi _(Trouessart), — Uromiys

caudimaculatus (Krefft), Zvzomys argurus (Thomas)

and Zyzomys woodwardi (Thomas); and Murinae,

*School of Biological and Environmental Sciences, Central

Queensland University, Rockhampton, Queensland 4700.

Australia. Email: Lwarner@cqu.edu.au

Rattus fuscipes, R. lutreolus (Gray) and R. sordidus

(see Smales, 1997). Species of Odilia have also been

found in the hydromyine Mallomys rothschildi

Thomas from Irian Jaya, now Papua, Indonesia and

the murines Rattus xanthurus Gray and Maxomys

musschenbreokii Jentink from Sulawesi, Indonesia

(Hasegawa & Sayaffrudin, 1994; 1995 ; Hasegawa et

al., 1999),

Rattus leucopus, the Cape York rat, and R.

sordidus, the canefield rat, are both found in

northern Queensland and the island of New Guinea,

the only two endemic murines that are found on both

sides of Torres Strait (Flannery, 1995). This

distribution provides evidence of recent past land

bridges, probably in the Pleistocene, between the two

land masses (Moore & Leung, 1995). Observations

on the host range of the species and the significance

of the geographical distribution of the genus Odilia,

given the geographical distributions of the hosts are

presented.

Materials And Methods

The specimens from R. leucopus were fixed in

10% formalin, stored in 70% ethanol and examined

in lactophenol. En face preparations and transverse

sections were cut by hand using a cataract scalpel

and mounted in polyvinyl — lactophenol.

Measurements of 10 males and 10 females from

Rattus leucopus were taken using an ocular

micrometer and given as the range followed by the

mean in parentheses, in micrometres unless

otherwise stated. Drawings were made with the aid

of an Olympus BH Nomarski interference contrast

microscope and drawing tube. Specimens are held in

the CSIRO Wildlife collection, Canberra (CSIRO)

and the South Australian Museum, Adelaide (SAM

AHC). Terminology and classification used follows

60 L. R. SMALES

Figs. 1-14. Odilia emanuelae (Mawson, 1961). Female head, en face view. 2. Female, anterior body section through

oesophageal region, arrow indicating axis of orientation of synlophe. 3. Male, mid body section. 4. Female, anterior

end, lateral view showing origins of ridges of synlophe. 5. Female, mid body section. 6. Male, section within posterior

third of body 7. Female, anterior end, lateral view. 8. Female, section within posterior quarter of body. 9. Genital cone,

right ventral view. 10. Bursa, dorsal view, flattened. 11. Spicule tips, lateral view. 12. Genital cone, right lateral view.

13. Female, posterior end left lateral view. 14. Male, posterior end, right lateral view. Abbreviations d, dorsal; |, left; r,

right; v, ventral. Scale bars: 1, 9, 11, 12, 10 um; 2, 3, 5, 6, 8, 20um; 4, 10, 25um; 7, 13, 14, 50m.

A REDESRIPTION OF ODILIA EMANUELAE 61

Durette-Desset (1971, 1973, 1983, 1985), Beveridge

& Durette-Desset (1992) and Durette-Desset ef al.

(1994). Rodent classification follows Strahan

(1995).

Odilia emanuelae (Mawson,1961)

(Figs 1-14)

Heligmonoides emanuelae Mawson, 1961, pp 809-

810, figs 30-34 table 4, from Rattus conatus (syn R.

sordidus ) and R, assimilis (syn R. fuscipes); Durette-

Desset (1969) p 738, fig 4C.

Austrostrongylus emanuelae: (Mawson,1961)

Durette-Desset 1971 p 65.

Odilia emanuelae: : (Mawson,1961) Durette-

Desset 1973 p 517; Smales 1992 p 75.

Material examined

From Ratius sordidus: holotype male, allotype

female, Innisfail, (17° 32’ S, 146° O1’ E) Queens-

land, SAM AHC 41332: from Rattus leucopus: 55

males, 81 females, East Mc Illwraith Range, Cape

York Peninsula (13° 45’ S, 143° 20’ E), Queensland

coll. P. Catling, I. Mason and P. Haycock, 9. xiii.

1990, 10. iti. 1990, CSIRO N3293, N3296, N3324,

N3325, N3326, N3329: from Melomys cervinipes

(Gould) 10 males, 10 females, D’Aguillar Range

(27° 50’ S, 152° 45’ E), Queensland, 19. ii. 1963,

SAM AHC 5805, coll. Aland and Stewart, 26.viii.

1993, SAM AHC 32190, 32191, 32192. Comparison

of the measurements of specimens from Rattus

sordidus, type host, and R. leucopus are given in

Table 1.

Redescription

Small coiled nematodes; prominent cephalic

vesicle present; buccal capsule vestigial. Mouth

opening triangular with rudimentary lips, surrounded

by four double papillae, each comprising a cephalic

plus externo-labial papilla and two lateral amphids.

Internal labial papillae not visible. Oesophagus

claviform; nerve ring surrounds oesophagus at about

mid level; excretory pore and digitiform deirids at

same level, posterior to nerve ring.

Synlophe: Longitudinal cuticular ridges

continuous, extend from posterior margin of cephalic

vesicle to just anterior to bursa or vulva; 17 in

anterior, 18 in mid body; axis of orientation from

right ventral to left dorsal at approximately 75° to

frontal axis; 7-8 in dorsal side, 9-10 in ventral side;

ridges | and 1’ largest, forming typical type A

carene, ridges 2, 3 smaller than ridges 1, ridges 4-6

increasing in size, ridges 7-10 decreasing in size.

Posterior region of body with 15 (male), 17 (female)

ridges reduced in size; dorsal side with 7-8; ventral

side with 7-10 ridges.

Male

Length 1.3-1.64 (1.50) mm, maximum width 54-67

(60). Cephalic vesicle 42-56 (50.6) long. Oesophagus

300-420 (345) long; excretory pore 231, 340 from

anterior end. Bursa asymmetrical, right lobe larger

(rays of right lobe more robust); deep dorsal cleft.

Dorsal ray symmetrical divided at about half its

length, each branch dividing again at distal tip;

terminal divisions, rays 9, 10 symmetrical; rays 8

arising at same level, right ray 8 more robust than left.

TABLE 1. Measurements of Odilia emanuelae, in {um unless otherwise stated, from two host species; 10 males and 10 females

from each. Data for Rattus fiscipes are from Mawson (1961).

Rattus leucopus

Rattus fuscipes

Locality East Mclllwraith Range Innisfail

Male

Length, mm 1.3-1.64 2.2-2.6

Width 54-67 -

Cephalic vesicle 42-56 50-60

Oesophagus length 300-420 240-270

Ant. end to excretory pore 230, 340 190-220

nerve ring - -

deirid - 190-220

Spicules 245-270 250-330

Gubernaculum 17-22 -

Female

Length, mm 1.8-2.6 2.1-3.1

Width 63-74

Cephalic vesicle 50-56 50-60

Oesophagus length 240-410 270-290

Ant. end to excretory pore - 200-222

nerve ring - -

deirid - 200-222

Tail 30-43 40-50

Eggs 50-63 x 27-34 60-80 x 40-50

Vulva to tail tip 87-119 105-130

62 L. R. SMALES

Rays 4, 5, 6 with common stem, reaching margin of

bursa; rays 4 and 5 robust curving anteriorly, rays 6

slender, curving posteriorly. Rays 2 and 3 with

common stem, robust, diverge distally, curve

posteriorly, reaching margin of bursa. Genital cone

short, ventral lobe with unpaired papilla 0, lightly

sclerotized; dorsal lip bifid, each lobe with single

papilla 7. Spicules equal, filiform, tips pointed, 245-

270 (253) long. Gubernaculum 17-22 (19.5) long.

Female

Length 1.8-2.6 (2.1) mm; maximum width 63-74

(67). Cephalic vesicle 49.5-56 (51.5) long;

oesophagus 240-410 (320) long. Vulva opens 87-119

(94.5) from tail tip; posterior end may or may not be

flexed at right angles just behind vulva.

Monodelphic, ovejector with sphincter 30, 35,

shorter than vestibule 50, 60, infundibulum, about

same length as sphincter. Tail 29.5-43 (35) long.

Eggs in utero 49.5-63 (56.5) by 26.5-34 (30).

Key to species of the genus Odilia

1. Synlophe with discontinuous ventral ridges........

Phat B hort dba O. mackerassae (Mawson, 1961)

Synlophe with continuous ridges..........:0cee 2

2. Synlophe with 18 or more ridges in mid body ..3

Synlophe with fewer than 18 ridges in mid body

PETTUS TRY Nt cin vir sneer 20 Canes crerceerrns 10

3. Synlophe with 18 ridges in mid body................ 4

Synlophe with more than 18 ridges in mid body

Heegesesaaesuei as atin bob pasa peated ese repeqedtnontuned shies 5

4, Synlophe with fewer than 18 ridges posteriorly,

12-15 in males. Gubernaculum 17-22 long;

spicule tips taper to sharp point. Female tail

conical, rounded tip ......... eee O. emanuelae

(Mawson, 1961)

Synlophe with more than 18 ridges posteriorly,

24 in males. Gubernaculum 30-40 long; spicule

tips joined distally surrounded by transparent

membrane. Female tail tapers sharply from vulva

to pointed tp... eee ee eee O. tasmaniensis

Gibbons & Spratt, 1995

5. Synlophe with 19-20 ridges (male) in mid body,

ridges becoming tiny posteriorly. Gubernaculum

19-22 long; spicule: body length 1: 6.5. Female

tail 30; eggs 59-78 x 29-42........ O. mamasaensis

Hasegawa, Miyata & Syafruddin, 1999

Synlophe with more than 20 ridges (male) in mid

BODY ccacsegeiy lst beg cecbaa crc ne tenn btn teesestaten 6

6. Synlophe with 21 ridges (male) in mid body.

Gubernaculum 28 long; spicule: body length 1: 8

pe caper eae O. mawsonae (Durette-Desset, 1969)

Synlophe with more than 21 ridges in mid body

7. Synlophe with up to 35 ridges in mid body ...... 8

Synlophe with more than 35 ridges in mid body

8. Synlophe with 24-25 (male), 24-28 (female)

ridges in mid body, 23 (male) 13 (female) ridges

becoming minute posteriorly. Gubernaculum 32-

37; spicule: body length 1:4. Female tail conical,

pointed tip; eggs 72-80 x 35-43 .....O. maxomyos

Hasegawa, Miyata & Syafruddin, 1999

Synlophe with 22-29 ridges (male), 24-35 ridges

(female) in mid body, 29 (male), 26 (female)

ridges posteriorly. Gubernaculum absent; spicule:

body length: 1:13. Female tail with prepuce; eggs

60-70 X AO bgcetescetetsrnenennrcetaneinne O. praeputialis

Gibbons & Spratt, 1995

9. Synlophe with many (male), 36 (female), even

sized ridges in mid body. Gubernaculum 20 long;

spicule: body length ratio 1:8. Female tail twisted

into 1-2 coils in front of vulva... eee

Jumepnaneatedarreaeeoel O. polyrhabdote (Mawson,1961)

Synlophe with 40 (male), 48 (female) ridges in

mid body. Spicule tips with hair like projection

60 from distal end supporting fan-like alae;

spicule: body length 1:9, Female tail conical 60-

70 long with prepuce .......... cece O. uromyos

(Mawson, 1961)

10. Synlophe with 17 ridges in mid body, 20 (male),

19 (female) posteriorly. Gubernaculum 50 long;

spicule: body length 1:14. Eggs 70-80 x 40-50

3 O. bainae Beveridge & Durette-Desset, 1992

Synlophe with fewer than 17 ridges in mid body

SAO OT NE ALT PLEO EITT fi aey ey reer OTEET ERED 11

11. Synlophe with 16 ridges (male) in mid body, 16

ridges smaller posteriorly. Spicule: body length

1:16. Female tail 50 long; eggs 69-77 x 35-45 ...

cates O. mallomyos Hasegawa & Syafruddin, 1994

Synlophe not as above ........cc cc eeeeeeeeereeeees 12

12. Synlophe with 15 (male), 16 (female) ridges in

PAA DOH aged betaatza rsd bacecactesatdossscanteneteerenss 13

Synlophe with less than 15 (male), 16 (female)

ridges in Mid DOY ....... eee eeeeeeer eee rreressesenees 14

13. Synlophe with 16-20 minute ridges posteriorly.

Gubernaculum 32-43 long; spicule: body length

1:7. Female cuticle inflated proximally to tail.....

tas O. moatensis Hasegawa & Syafruddin, 1999

Synlophe with 30 (male), 50 (female) small even

ridges posteriorly. Gubernaculum 51-62 long;

spicule: body length 1:8. Female cuticle not

inflated proximally to tail.......... O. sulawesiensis

Hasegawa & Syafruddin, 1999

14. Synlophe with 14 (male), [5 (female) ridges in

mid body, 16 (female), smaller even ridges

posteriorly. Gubernaculum 30 long; spicule tips

pointed; spicule: body length 1: 9-1: 11. Female

tail conical, rounded tip 40-50 long wu...

be ates actin hs ony eg O. melomyos (Mawson,1961)

Synlophe with 14 ridges (male) in mid body.

Gubernaculum 25 long; spicule tips expanding,

bifid; spicule: body length 1:9. Female tail

conical, pointed tip, flexed sharply back on itself

Babine dete coadahs deg ME O. brachybursa (Mawson,1961)

A REDESRIPTION OF ODILIA EMANUELAE 63

Although Hasegawa & Sayafruddin (1995) noted

Odilia sp. 1 and Odilia sp. 2 from Rattus cf.

morotaiensis — from Indonesia, insufficient

morphological data were provided to allow inclusion

of these two species in the key.

Discussion

The present study of O. emanuelae revealed slight

variations in the morphology of the species as

compared with the description of Mawson (1961),

particularly in the range of measurements, with

specimens from R. leucopus smaller than those from

R. sordidus. The spicule tips were described as

widened and alate by Mawson (1961). Gibbons &

Spratt (1995), however, commented that they

broadened then tapered to a sharp tip. Examination

of the type male, as well as the specimens from 2.

leucopus in this study confirmed this latter form. The

ridges of the synlophe were counted by Mawson

(1961) as up to 20 in the mid body but given and

figured as 18 by Durette-Desset (1969) for a female

worm from R. sordidus, as is the case for specimens

from R. leucopus. These minor morphometric

differences may be due to host induced variation and

are not sufficient to establish a separate species.

Consequently the material from R. leucopus is

assigned to O. emanuelae. The host range is

accordingly expanded to include a third endemic

Rattus species.

The only other host records for O. emanuelae are

from M. cervinipes, specimens from a single host

deposited in the SAM and specimens collected

during an unpublished survey of the helminths of M.

cervinipes and R. fuscipes from the D’ Aguilar

Ranges, south east Queensland (Aland!) in which

two of 12 M. cervinipes were reported as being

infected with O. emanuelae. Re-examination of this

material revealed specimens of O. emanuelae in a

third host, making a total of 4 infected M. cervinipes,

all from the D’Aguilar Ranges. Odilia emanuelae

has not been reported in other surveys of the

helminths from melomys, such as that of Mawson

(1961) although she examined hosts from Innisfail,

the type locality (Mawson, 1961; Smales, 1997).

This suggests that Raffus species are the normal

hosts and that infections found in M. cervinipes in

' Aland, K. (1993) BSc Hons Thesis, Dept of Parasitology,

University of Queensland.

this study are an example of an occasional infection

occurring where normal and alternative hosts are

sympatric. The geographic distribution of O.

emanuelae has been extended further north from the

type locality into Cape York and south to south east

Queensland.

Previous interpretations of the origins of the

Trichostrongylina in Australian rodents have

presumed that the genus Odilia arose in Australia,

co-evolving with the rodent sub family Hydromyinae

(see Durette-Desset, 1985); that is with the earliest of

the rodent invaders commonly known as the old

endemics. The rodents are thought to have arrived in

Australia some 5-10 million years ago (Watts &

Aslin, 1981; Flannery, 1995). Odilia was then

captured by more recent rodent arrivals, the Murinae

new endemic Rattus species that crossed to Australia

from New Guinea less than one million years ago

(Smales, 1992; Beveridge & Durette-Desset, 1992;

Gibbons & Spratt, 1995),

More recently, however Odilia species have been

described from several Indonesian islands and

occurring in both old endemic and new endemic

hosts (Hasegawa & Syafruddin 1994; 1995;

Hasegawa ef al., 1999). At the same time, new fossil

evidence from Australia suggests more complex

evolutionary processes than had first been thought.

There is now evidence for at least three phases of

immigration, both direct from Southeast Asia, and

through New Guinea, involving both old and new

endemics (Godthelp, 2001).

This new evidence suggests that ancestral forms of

Odilia may have co-evolved with rodent hosts in

Southeast Asia. The present host and geographic

distribution therefore reflects a complex series of

evolutionary events involving host switching and co-

evolution as rodent faunas and their helminth

communities migrate, and undergo evolutionary

radiations, The more data gathered about rodent

hosts and their parasites from Southeast Asia and

Australasia the more complex the patterns of their

relationships become.

Acknowledgements

I am grateful to Dr D. Spratt for making the

material available.

64 L. R. SMALES

References

BeveERIDGE, I. & DurRETTE-DEsseT, M.-C. (1992) A new

species of trichostrongyloid nematode, Odilia bainae,

from a native rodent, Rattus fuscipes (Waterhouse).

Trans. R. Soc. S. Aust. 116, 123-128.

DuretTtE-Dessert, M.-C. (1969) Les systemes d’arétes

cuticulaires chez les Nematodes H¢éligmosomes parasites

de muridés Australiens. Ann. Parasitol. Hum. Comp. 44,

733-747.

(1971) Essai de classification des Nématodes

Héligmosomes. Corrélations avec la paléobiogéographie

des hétes. Mém. Mus. natn. Hist. nat. Série A, Zoologie,

69, 1-126.

(1973) Note réctificative sur le genre

Austrostrongylus (Nématode). Ann. Parasitol. Hum.

Comp. 48, 517.

(1983) Keys to the genera of the superfamily

Trichostrongyloidea, No. 10 pp.1-86 /n Anderson R. C.

& Chabaud A. G. (Eds), “CIH Keys To The Nematode

Parasites Of Vertebrates ” (Commonwealth Agricultural

Bureaux, Farnham Royal).

(1985) Trichostrongyloid nematodes and their

vertebrate hosts: reconstruction of the phylogeny of a

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

, BeveripGE, I. & Spratt, D. M. (1994) The

origins and evolutionary expansion of the Strongylida

(Nematoda). Int. J. Parasitol. 24, 1139-1165.

FLANNERY, T. (1995) “Mammals of New Guinea” (Reed

Books, Chatswood).

Gipsons, L. M. & Spratt, D. M. (1995) Two new species of

Odilia (Nematoda: Heligmonellidae) from Australian

rodents, with comments on O. bainae Beveridge &

Durette-Desset 1992. Syst. Parasitol. 31, 67-79.

GopTHELp, H. (2001) The Australian rodent fauna, flotilla’s

flotsam or just fleet footed pp. 319-321 /n Metcalfe, 1.

Smith, J.M.B. Morwood, M. & Davidson, I. (Eds)

“Faunal and Floral Migrations and Evolution in S.E. Asia

— Australasia” (A.A. Balkema, Lisse).

Hasecawa, H. (1996) Notes on the morphology of three

nematode species of the subfamily Nippostrongylinae

(Heligmosomoidea: Heligmonellidae) collected from an

endemic rat of Halmahela Island, Indonesia. Biol. Mag.

Okinawa, 34, 13-21.

& SAYAFRUDDIN, (1994) Odilia mallomyos

sp.n. (Nematoda: Heligmonellidae) from Mallomys

rothschildi weylandi (Rodentia: Muridae) of Irian Jaya,

Indonesia. . Helm. Soc. Wash. 61, 208-214.

(1995) Nippostrongylus

marhaeniae sp. n. and other nematodes collected from

Rattus cf morotaiensis in North Halmahera, Molucca

Islands, Indonesia. /bid. 62, 111-116.

, Miyata, A. & SYAFRUDDIN, (1999) Six new

nematodes of the Heligmonellidae (Trichostrongylina)

collected from endemic murines of Sulawesi, Indonesia.

J. Parasitol. 85, 513-524.

Mawson, P. M. (1961) Trichostrongyles from rodents in

Queensland, with comments on the genus Longistriata

(Nematoda: Heligmosomatidae). Aust. J. Zool. 9, 791-

826.

Moorg, L. A. & LEUNG, L. (1995) Cape York Rat Rattus

leucopus (Gray, 1867) pp. 635-655 Jn Strahan, R. (Ed.)

“The Mammals of Australia” (Reed Books, Chatswood).

SMat_es, L. R. (1992) A survey of the helminths of Rattus

sordidus with a description of Ancistronema coronatum

n.g., n.sp. (Nematoda: Chabertiidae). Syst. Parasitol.

22, 73-80.

(1997) A review of the helminth parasites of

Australian rodents. Aust. J. Zool. 45, 505-521.

STRAHAN, R, (1995) “The Mammals of Australia” (Reed

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Transactions of the Royal Society of S. Aust. (2005), 129(1), 65-73.

THREE DECADES OF HABITAT CHANGE IN GULF ST VINCENT,

SOUTH AUSTRALIA.

by J. E. TANNER"

Summary

TANNER, J. E. (2005). Three decades of habitat change in Gulf St Vincent, South Australia. Trans. R. Soc. S. Aust.

129(1), 65-73, 31 May, 2005.

Benthic habitats in Gulf St Vincent, South Australia, have changed considerably in the period between the

1960s and 2000/2001. Diver surveys in the 1960s indicated the presence of an extensive area of deep-water

Heterozostera seagrass in Investigator Strait, and Ma/leus-Pinna bivalve assemblage in the south-eastern area of

the gulf. Neither of these habitat types were detected in remote video and diver surveys conducted in 2000/2001,

with these areas instead being dominated by relatively barren sand flats. In the central section of the gulf, the

cover of bryozoans, and density of scallops, also declined over the three decades between the two surveys.

Further north there have been fewer changes, with the original Pinna assemblages and seagrass meadows still

present. It is thought that these changes are primarily due to anthropogenic influences, with the two major

candidates being increased turbidity (due to terrestrial inputs from sewage, stormwater runoff, agricultural runoff

and dust storms), and direct damage from prawn trawling. As a consequence of these changes, the habitat

complexity in Gulf St Vincent has decreased substantially, with likely consequences for other fauna such as fish

and mobile invertebrates.

Introduction

While the effects of anthropogenic disturbance on

many terrestrial systems are obvious and well

documented, much less is known about what changes

are occurring in marine systems, especially in waters

deeper than a few metres. It is often thought that

marine systems are stable, and little affected by

change on a broad-scale, despite some well

documented examples of change due to specific

disturbances at smaller scales (e.g. Trawling:

Jennings & Kaiser 1998; Hall 1999; Pollution:

Neverauskas 1987; Lapointe et al, 1994; Sea Level

Change: Seddon ef a/. 2000). This attitude is

probably related to our inability to easily see what is

happening on the ocean floor, and to the lack of long-

term studies and baseline data on what systems

looked like before human disturbance (or even in its

early stages). While there is good evidence of change

in individual habitats in some marine systems, such

as seagrass loss (e.g. Neverauskas 1987; Walker &

McComb 1992; Short & Wyllie-Echeverria 1996;

Edyvane 1999), and changes in kelp abundance in

the eastern North Pacific related to changes in otter

abundance (e.g. Estes & Duggins 1995), there are

few well documented studies that examine change in

entire marine ecosystems over a period of decades.

Most of those studies that have been done rely on the

existence of earlier studies that utilised remote

sampling techniques (such as benthic grabs) to

examine infauna, or trawl gear to examine catches of

" SARDI Aquatic Sciences, PO Box 120, Henley Beach, SA 5022.

Australia. Phone: +61 8 8207 5489. Fax: +61 8 8207 5481,

Email: tanner,jason@saugoy,sa.gov.au.

fish and other macrofauna (e.g. Haedrich & Barnes

1997; Wilson et al. 1998; Frid et al. 1999, 2000), and

have not specifically examined changes in benthic

habitats.

Between 1964 and 1969, Shepherd & Sprigg

(1976) conducted an extensive series of diver surveys

of benthic habitats in Gulf St Vincent, South

Australia. They observed the intact habitat in situ,

and recorded all the major components of the flora

and fauna, While these surveys were primarily

qualitative, with few abundance estimates, a

comprehensive map documenting the various

community types in the gulf was _ published

(reprinted here as Fig. 1). This map was used to

compare the major habitat features and benthic

assemblages present in the 1960s, to those present in

2000/2001, to detect any substantial changes in the

intervening period, To do this an extensive series of

remote video surveys was undertaken in 2000/2001,

complemented by a number of spot dives to ensure

that the information being obtained from the video

was reliable. There have been no extensive benthic

surveys carried out in Gulf St Vincent between these

two studies, and thus we do not know if any

intermediate states occurred, and it is only possible

to speculate on the causes of any changes observed.

Gulf St Vincent is a large (~ 13000 km? including

its approaches), relatively shallow (maximum depth

~ 40 m), marine embayment located on the South

Australian coast. It is an inverse estuary, with salinity

ranging from ~37%. at the mouth to 41%. at the head,

due to high evaporation rates and low precipitation

(Bye 1976). The location of Kangaroo Island across

the mouth of the gulf means that exchange of water

with the open ocean is restricted, leading to long

66 J. E. TANNER

137°30'E 138°0'E 138°30'E

Bare sand and shoal

Heterozostera-Lunulites

assemblage

Algal debris (Calcarenite)

mostly desert

Ascidian-scallop

assemblage

Pinna-holothurian

assemblage

34°30'S

Bryozoan assemblage

34°30'S

Malleus-Pinna

assemblage

Seagrass meadows.

Mostly Posidonia

Boulder conglomerates

Reef (kunkerised shell

beds) of low relief

TARE Oi

Aeolianate reef

2000 survey point

35°0'S

=== ee ee ee et

xxx x x

xx KK xX

XXXXXK

cK MK XK OK RK OL KK XK

J Kak x Wah ahah oh x% « & x OS

MM RM MM RM MM MRK M KKK KRM KM KK KO!

35°30'S

137°30'E 138°0'E 138°30'E

Fig. 1. Benthic community composition in Gulf St Vincent in the 1960s as determined by diver spot surveys (modified from

Shepherd and Sprigg 1976). The survey sites for 2000/2001 are marked for comparison.

THREE DECADES OF HABITAT CHANGE 67

residence times of water within the gulf (de Silva

Samarasinghe & Lennon 1987). This geography also

means that the area is a low energy environment,

especially in the northern section, which as a

consequence, is slowly being filled in by sediment

deposition. Most of the substrate is either sand or fine

silt, with only a few areas of hard bottom (Shepherd

& Sprigg 1976, see Fig. 1), although there are

substantial areas of calcrete underlying much of the

sand, The city of Adelaide (population ~ | million), is

located on the eastern shore of the gulf, and is a

source of domestic and industrial pollution. The only

other settlements in the area are several small towns

with populations < 1000. There is considerable

agricultural activity along the shores of the gulf, and

in its catchment, which is a further source of

pollution, However, due to the arid nature of the

region (precipitation < 500 mm yr") there is little

natural runoff. The gulf also supports substantial

recreational and commercial fisheries, including a

small (10 boat) prawn fishing fleet that targets the

western king prawn (Melicertus latisulcatus).

The prawn fishery commenced in 1968/69, and

effort reached a peak of 15200 hours of trawling in

1982/83, and a peak catch of 602 t in 1976/77, before

dramatic declines in both catch (to 200-400 t yr!)

and effort (to ~4000 hours yr’) for the period 1995-

2002 (Svane & Johnson 2003). During the early

phase of the fishery, the mid-northern portion of the

gulf was heavily targeted (but south of 34° 30’ S), but

since the mid 1980s to early 1990s effort has

predominantly focussed on more southern areas

where the prawns tend to be larger (Morgan 1995).

Materials and Methods

To quantify the distribution, abundance and

composition of benthic habitats throughout Gulf St

Vincent, an extensive series of remote video surveys

was conducted between June 2000 and June 2001.

Survey sites were located every 2 nm along east-west

transects across the gulf which were spaced every 5

nm (18 transects between 34° 15’ S and 35° 40’ S).

Transects extended from the eastern to the western

edge of the gulf (although areas less than

approximately 5 m deep were inaccessible to the

vessel used and therefore excluded). The western

extremity of the survey area was 137° 40’ E, and the

easternmost point was 138° 30’ E. In total, data were

obtained for 294 sites, with an additional three sites

excluded from analysis because of poor image

quality (see Fig. 2 for site locations). At each site, a

digital video camera was lowered to within | m of

the bottom, and left to record for 10 min while the

boat drifted. To determine the linear distance moved

during this time, a GPS was used to record the

location (+10m) when a clear image of the bottom

was first obtained, and again when it was lifted off

the bottom. The mean distance covered per 10 min

survey was 141+] m (se).

Animal taxa visible in the video footage were

enumerated, with total abundance standardised to the

mean distance covered in a 10 min survey (141 m).

Counts were only made for segments of the footage

that were clear, and standardised counts were also

adjusted for the proportion of the video that could

not be used reliably. The exact area covered could not

be calculated, as there was some variation in the

height of the camera from the substrate, however, on

average a swathe of ~ 2 m wide was surveyed. For

those taxa for which individuals could not be

distinguished (seagrasses and algae), as well as bare

substrate, the video was stopped during playback

approximately every 1 min, and percent cover

recorded with the aid of a grid overlying the image.

The mean percent cover for all recorded frames was

then calculated for each site. To ensure that the video

provided a reliable record of the benthos, spot dives

were made at 53 sites to examine the benthos in more

detail, and a series of photographs were taken of 0.25

m? quadrats for a later comparison with the video

footage. These photographs were only qualitatively

assessed, and did not show any major discrepancies

with the video footage, so are not considered further.

Dives were conducted on most of the defined habitat

types, although depth constraints prevented stations

> 24 m in depth from being surveyed in this way.

To objectively determine the community type at

each site, cluster analysis was used on the

standardised data. As the objective was to produce a

map that could be compared with that produced by

Shepherd & Sprigg (1976), taxa were grouped at the

same level as they used (e.g. sponges, bryozoans,

scallops etc), and minor taxa were removed from the

analysis. The clustering technique used was Wards

Flexible B, with B = -0.25 (Seber 1984). This

technique was chosen as it did not produce any

chaining, unlike more commonly used methods such

as group-average and centroid. Initially, 15 groups

were chosen for further investigation, and these were

manually merged on the basis of their dominant taxa

to achieve similar groupings to those used by

Shepherd & Sprigg (1976). Not all of the habitat

types defined by Shepherd & Sprigg (1976) were

present in the 2000/2001 survey, while several new

intermediate habitat types were defined based on

small clusters that did not readily fit in with any of

the original types. It should be emphasised that the

2000/2001 survey sites did not correspond to the

survey sites used in the 1960s, as information on the

location of the later was not available. Thus, any

comparisons over time are based on_ the

interpolations made by Shepherd & Sprigg (1976) in

producing their map of habitat distributions.

68 J. E. TANNER

137°30'E 138°0'E 138°30'E

Legend

Barren Sand

Ascidian

Ascidian/Bryozoan

34°30'S

Bryozoan fe +XxX ofm 7

a =~ “)

inna a+xcox@f snnas

{\ oa. 2, eer

34°30'S

Scallops

a eee

. Se,

} |

Seagrass/Pinna Z Wy ae a

P+ + mot x eee

Seagrass (| |

fan docxrxs ences ane

Aomexr xxop+exkmeoend

AW\| /

maxx me ee eae ee

\ 7,

— Pie Le SES tSbe!

Pe ereterren Sree rre rey

/\,

Ais eae

35°0'S

xx xX XX XX KKK KKXKXXK XX

35°30'S

ee ne het

(

ee, eS —L

35°30'S

a \ ‘ _

an mxxx xX XXX xXx

s'm XX XE EXX

2S is.

137°30'E 138°0'E 138°30'E

Fig. 2. Benthic community composition in Gulf St Vincent in 2000/2001 as determined from remote video surveys.

outlines of the habitats present in the 1960s are included for comparison.

The

THREE DECADES OF HABITAT CHANGE

Results

There have been some substantial changes in the

epibenthos of Gulf St Vincent since the surveys of

the mid to late 1960s by Shepherd & Sprigg (1976)

(compare Fig. | with Fig. 2, Table 1). Particularly

noticeable is the absence of the seagrass

Heterozostera tasmanica, which covered extensive

areas of deep sand plains in the southern gulf and

Investigator Strait in the 1960s. While there was only

a sparse cover of Heterozostera in the 1960s (S.

Shepherd pers. com.), this area is now completely

devoid of seagrass, with none being seen either in the

remote video footage or on the spot dives. Also

missing is the Malleus-Pinna (bivalve) assemblage

that Shepherd & Sprigg (1976) documented in the

south-eastern section of the gulf. While this area still

contains some scattered Pinna (<0.1 m~), there was

no evidence of any Malleus (hammer oysters that

grow up to 150 mm in length). It is possible in this

case that individual animals would not have been

detected in the remote video footage because of their

cryptic nature, whilst the depth precluded diving to

check for them, Nevertheless, the clumps that existed

previously would have been detectable with the video

sampling, and can thus be regarded as absent. Both

of these assemblages are characterised by relatively

long-lived organisms (5-10 yrs), and thus short-term

seasonal or annual variation is unlikely to explain

their absence in 2000/2001.

There is no evidence that new types of assemblage

dominated by large macrofauna/flora have

established in the place of the Heterozostera and

Malleus-Pinna assemblages that have disappeared.

Instead, these areas now appear to be predominantly

bare sand, with scattered invertebrates, including

ascidians, bryozoans, sponges and some Pinna

(Table 2). There are, however, extensive areas

(mostly in water greater than 30 m deep) that are

very depauperate in large macrofauna, and appear to

be barren sand plains (Fig. 2), at least with respect to

epibenthic organisms.

The other substantial changes are an apparent 80%

decrease in the area dominated by bryozoans in the

central part of the gulf, and a reduction in the

abundance of scallops in the central eastern section.

In most of the areas where scallops were formerly a

noticeable part of the benthic assemblage (with

densities of 0.5 — 4 m*, Shepherd & Sprigg 1976)

they now only occur in very low abundance (Table

2). In 2000/2001, only three sites on the western side

of the gulf, and one in the south-east, had substantial

TABLE 1. Habitat change, measured as the percentage of each habitat type that changed to other habitat types, in Gulf St

Vincent over the period 1964-69 to 2000-01, Left column represents habitats in the 1960s, top row habitats in 2000. It

should be noted that the habitat present in the 1960s was based on interpolation from dive sites that did not correspond

to the 2000/2001 survey sites, potentially inflating the differences between the two surveys.

Barren Ascidian Ascidian Bryozoan Pinna Scallop Seagrass Seagrass Total

Sand Bryozoan Pinna transitions

Heterozostera 100% 55

Algal debris 50% 50% 2

Ascidian-scallop 29% 47% 3% 3% 13% 38

Pinna-holothurian 31% 6% 6% 56% 5% 16

Bryozoan 56% 2% 21% 21% 43

Malleus-Pinna 95% 5% 42

Seagrass 15% I% 5% 1% 10% 3% 3% 62% 78

Reef (kunkerised shell) 33% 33% 33%

Aeolianate reef 100% 2

TABLE 2. Densities and percent cover of major taxa in each habitat category in 2000/2001. Habitat categories are based

on a cluster analysis, as described in the text. Density values assume each transect is 2 m wide, and as transect width

could not be measured, are only approximate. Numbers in parentheses are standard errors. Blank cells indicate that the

taxon did not occur in that habitat.

Ascidian Pinna Bryozoan Sponge Scallop Sand Algae Seagrass

(%) (%) (%)

Ascidian 2.2 (0.5) 0.07 (0.03) 0.01 (0.008) 0.8 (0.7) 0.05 (0.02) 89 (3) 6 (2) 4 (2)

Ascidian — Bryozoan 3.6 (0.04) 0.05 (0.03) 3.6 (0.04) 0.8 (0.7) 0.07 (0.05) 73 (8) 2 (1) 13 (5)

Bryozoan 0.05 (0.03) 0.3 (0.3) 4.1 (0.3) 0.2 (0.04) 0.007 (0.004) 93 (2) 1 (1)

Pinna 0.06 (0.02) 0.9 (0.3) 0.09 (0.02) 0.5 (0.2) 0.003 (0.002) 84 (4) 8 (3) 5 (2)

Barren Sand 0.01 (0.002) 0.02 (0.002) 0.008 (0.001) 0.2 (0.05) 0.003 (0.001) 98 (0.3) 0.4 (0.1) 0.2 (0.1)

Scallops 0.05 (0.02) 0.06 (0.03) 0.9 (0.9) 0.07 (0.02) 3.7(0.08) 71 (22) 7(4)~——s«<717)

Seagrass 0.02 (0.005) 0.03 (0.01) 0.04 (0.008) 0.01 (0.009) 24 (4) 4(1) 72 (4)

Seagrass - Pinna 0.4 (0.3) 0.8 (0.3) 0.05 (0.004) 0.2 (0.1) 21 (2) 2 (1) 73 (6)

70 J. E. TANNER

numbers of scallops (Fig. 2), whereas in the 1960s

there were also large areas on the eastern side where

they dominated (Fig. 1). This reduction cannot be

attributed to commercial scallop fishing as no such

fishery has operated in the area. Bryozoans now only

occur in low abundance, mostly in the central part of

the gulf. Only 12 of the 294 sites surveyed had a

bryozoan cover of greater than 5%, with none greater

than 25%. This compares with the 1960s, when

bryozoans dominated a substantial portion of the

central gulf (Fig. 1, Table 1).

As was the case in the 1960s, seagrasses still

dominate many of the shallower areas <10 — 12 m

deep both in the gulf and along the northern shore of

Kangaroo Island (Fig. 2), with 65% of the sites

originally classified as seagrass still being either

seagrass or seagrass/Pinna in 2000/2001 (Table 1).

The dominant seagrass genus remains Posidonia,

with only a few areas of Amphibolis, and Halophila

occurring predominantly on the western side of the

gulf (see Tanner 2002). There is also some Halophila

in deeper waters, which appears to be ephemeral as it

was only detected at sites surveyed during the

summer. Pinna in the northern section of the gulf

remained relatively unchanged, although only 56%

(9 of 16) of sites originally classified as

Pinna/holothurian remained as Pinna, while a

further 24 sites changed from some other habitat to

Pinna (Table 1).

Discussion

There have been some obvious changes in the

benthic assemblages present in Gulf St Vincent in the

period between this study and that of Shepherd and

Sprigg (1976), especially in the southern part of the

gulf and in Investigator Strait. The main changes are

the loss of extensive deep-water Heferozostera

tasmanica meadows and Malleus-Pinna assemblages

in the southern region, and a reduction in the cover of

bryozoans and density of scallops in the central and

eastern parts of the gulf. Interestingly, it is the deeper

regions further from land that are generally the most

changed, although the very shallow (<5 m deep)

inshore areas were not included in this survey. There

are several significant anthropogenic influences that

may have contributed to these changes, although

natural processes may also have played a role. The

city of Adelaide is responsible for discharging a large

amount of pollution into the gulf, in addition to that

coming from agricultural runoff, and this is likely to

have imposed a substantial stress on many organisms

(Miller 1982; Neverauskas 1987; Edyvane 1999),

There were also several severe dust storms in the

1980s that removed large amounts of topsoil from

Yorke Peninsula (on the western shore) and deposited

it into gulf waters (G. K. Jones, pers. comm.). The

long flushing time of the gulf (Bye 1976; de Silva

Samarsinghe & Lennon 1987) will have exacerbated

any effects due to increased terrestrial inputs, as they

are only slowly removed from the system. Prawn

trawling has also been extensive (Morgan 1995), and

although now carried out in a much more sustainable

manner, damage from the previously intensive

fishery may take many decades to be reversed.

The loss of Heterozostera is most likely a result of

increased water turbidity, and a subsequent decline in

the amount of light reaching the bottom, although

unfortunately no historical turbidity data could be

found to determine the extent of this change. Given

that this species occurred predominantly in deep (30-

40 m) water, it was probably at its lower depth limit,

and it would only have required a small decrease in

light penetration for Heterozostera to be unable to

maintain itself. The maximum recorded depth for

this species is 39 m (Duarte 1991). An increase in

turbidity could have come about through several

different mechanisms, Firstly, increased coastal

discharge, both from the city of Adelaide and from

agricultural areas, may have resulted in an increase in

the amount of fine sediment in the water column, It

would thus be of interest to examine the sediments of

these and other areas of the gulf to see if an increase

in the amount of terrigenous material can be detected

over the last several decades. Secondly, there has

been a substantial loss of seagrass along the

metropolitan coast (~ 5000 ha), possibly connected

to sewage discharge (Neverauskas 1987; Shepherd er

al. 1989), resulting in a substantial increase in the

rate of sediment resuspension in shallow waters.

Although this increase in sediment resuspension has

not been reliably quantified, there is now a consistent

band of dirty brown water inshore of the seagrass

line, which numerous anecdotal reports suggest is a

relatively recent phenomenon (occurring since the

loss of seagrasses over the last 40-50 years). If these

resuspended sediments include a_ substantial

proportion of very fine material that can stay in

suspension for long periods of time, it is possible that

they may have been distributed throughout the gulf,

resulting in a system-wide increase in turbidity.

Finally, heavy trawling activity is well known to

result in sediment resuspension (Churchill 1989;

Pilskaln et a/. 1998; Palanques et a/. 2001), and this

may have increased the amount of suspended

material. Trawling may also have had a direct impact

on Heterozostera, causing more damage than could

be sustained in areas where it could only just survive.

The area formerly covered by Heferozostera has

experienced substantial trawling pressure throughout

the lifetime of the prawn fishery (Morgan 1995), and

despite the substantial decline in effort over the last

decade, the system may have experienced a state

change that cannot easily be reversed.

THREE DECADES OF HABITAT CHANGE 71

The former Ma/leus-Pinna assemblage that existed

in the south-eastern portion of the gulf coincides very

closely with the current main trawl grounds for the

prawn fishery. It is thus likely that this assemblage

experienced substantial direct damage from the trawl

gear, as well as possible negative effects from

increases in turbidity, whether due to trawling or

coastal activities. While most of this area could not be

examined by divers because of the depth, no Malleus

were seen either in the video footage, or in the

shallower areas that were accessible to divers. Further

north, the bryozoan assemblages would also have

been susceptible to trawl activity, as they are fragile

and not adapted to cope with extensive physical

disturbance (Bradstock & Gordon 1983). At its peak,

the prawn fishery trawled in excess of 1600 km? yr'!

(assuming a trawl speed of 3 kn, that the nets sweep a

20 m wide path, and that no areas are trawled more

than once ina year), and thus would have had a direct

effect on a large proportion of the gulf, although

currently a much smaller area is trawled (200-450 km?

yr! in the 1990s, J. Tanner unpublished data), While

there are many well documented cases of trawling

having a negative impact on benthic organisms (e.g.

Auster ef al. 1996; Engel & Kvitek 1998: Kaiser et al.

1998; Collie et al. 2000), there is considerable

controversy over how great the real impact is. This

controversy arises as many studies have failed to show

that trawling affects the benthic community (e.g.

Gibbs et al, 1980; Van Dolah ef al. 1991; Hall et al.

1993; Hannson ef al. 2000; Lindegarth et a/. 2000). In

Gulf St Vincent, the current trawling practices seem to

have little impact on infauna over the short term, at

least in the ascidian and Pinna habitats that have been

studied (Drabsch et a/. 2001), although approximately

36% of epifauna is removed or dies subsequently

(Tanner 2003). This suggests that previous periods of

intense trawling may well have had a substantial

negative effect on benthic communities in the gulf.

The loss of these macro-faunal and floral

assemblages has potentially important ecosystem

level consequences. As the species that have been

lost are those that formed most of the structure in

these otherwise relatively homogenous sand plains,

there are likely to be important implications for

species that require complex habitat-structure to

survive. For example, in New Zealand, bryozoan

beds form an important habitat for juvenile snapper

and other commercially fished species and have thus

been protected from destructive fishing practices

(Bradstock & Gordon 1983). Given that other

habitat-forming species have not replaced those that

have been lost, the physical complexity of these

habitats has been greatly reduced, and concomitant

losses or reductions in other species are likely. It is

well established, for example, that shallow-water

seagrasses provide important habitats for many fish

and invertebrate species (e.g. Bell & Westoby 1986;

Edgar 1990; Connolly 1994; Perkins-Visser et al.

1996), and while much less is known about deep-

water seagrasses, they are likely to serve a similar

function. These habitat-forming species may have

also provided an important food source for other

species, and so their loss may have disrupted food-

webs. For example, snapper (Pagrus auratus) are

important predators of Malleus, and the loss of the

latter species may have played a role in the decline of

snapper stocks in Gulf St Vincent.

In conclusion, there have been substantial changes

to the benthic habitats present in Gulf St Vincent

between the 1960s and 2000/2001, especially in the

central and southern regions. Potential causes

include increases in turbidity due to terrestrial runoff

(including sewage discharge, stormwater and

agricultural runoff) and dust-storms, and the direct

effects of trawling. The northern (most inland)

portion of the gulf seems to have experienced the

least degradation, possibly due to the very limited

terrestrial runoff associated with South Australia’s

arid climate, and the lack of a history of trawling.

The documented changes predominantly involve the

loss of important structure-forming species, and thus

could potentially have substantial implications for

many associated species that rely on physically

complex habitats for their survival.

Acknowledgments

I would like to thank T. Fowler, K. Jones,

S. Shepherd and two anonymous reviewers for

comments on an earlier version of this manuscript.

Thanks also to the many people who helped with the

surveys. Funding for this research was provided by

FRDC grant # 1998/208.

72 J. E. TANNER

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Transactions of the Royal Society of S. Aust. (2005), 129(1), 74-80.

FLOOD REGIME CHANGE IN THE HATTAH LAKES VICTORIA

RESULTING FROM REGULATION OF THE RIVER MURRAY

by N. J. SourER"

Summary

Souter, N. J. (2005) Flood regime change in the Hattah Lakes Victoria resulting from regulation of the River

Murray. Trans. R. Soc. S. Aust. 129(1), 74-80, 31 May, 2005.

Regulation of the River Murray was found to alter the frequency and variability of flooding through the Hattah

Lakes on the river’s floodplain. The height of Lake Hattah, recorded monthly since May 1908, was correlated

with historical data recording the extent of flooding through the system. This allowed the flooding frequencies

for six lakes in the system to be calculated using the remaining height data for Lake Hattah. An investigation of

pre (1908-1935) and post (1936-1996) regulation frequencies revealed a reduction in the frequency of small and

medium sized floods and a reduction in overall variability of flooding frequency throughout the system. Inflow

channel remodelling undertaken in 1972/73, in an attempt to increase the flooding frequency of the lakes,

increased the frequency of small floods similar to pre-regulation levels but did little to improve the variability

of flooding through the system.

Key Woros: River regulation, floodplain lakes, flooding frequency, environmental flows,

Introduction

Flooding frequency and the length of the dry

period between flooding are important factors in

structuring the aquatic communities of wetlands

(Crome 1986; Boulton & Lloyd 1992; Wellborn et al.

1996; Sheldon & Puckridge 1998). Differences in

flooding frequency have been related to differences

in macroinvertebrate diversity between floodplain

lakes (Van den Brink et al. 1994) and are known to

effect the emergence of aquatic invertebrates from

dry floodplain sediments (Boulton & Lloyd 1992).

Fish are known to utilise floodplains as habitat, as

adults (Welcome 1985; Reimer 1991) and as

nurseries for native Australian fish (Gehrke 1992).

Changes in vegetation structure may also occur as a

result of changes in flooding frequency (Bren 1992;

Roberts & Marston 2000; Roberts et a/. 2000). River

regulation through the construction and operation of

dams, weirs and locks to provide predictable flows

for human use, changes the natural flow regime of

* Floodplain Ecology Group, Department of Natural Resources and

Environment, Kaiela Research Station, Shepparton Victoria.

Current address: Department of Water, Land and Biodiversity

Conservation, Surface Water Assessment Branch, GPO Box 2834,

Adelaide SA 5001. Email: nsouter@adam.com.au

Siepentritt, M. & WILKINSON, L. 2004 The Murray-Darling

Basin’s Wetlands: Managing the Natural and Economic Riches

(Department of the Environment and Heritage, Canberra).

? BAKER, T. (1976) Report on investigations into the Hattah Lakes

National Park water supply problem. National Parks Service,

Victoria (unpub. ).

Suaw, J. R. (1985) The effects of river regulation and management

of water supplies on the Hattah-Kulkyne National Park and

Murray-Kulkyne park, Masters Thesis, University of Melbourne

(unpub.).

CummMING, P. L. F., & LLtoyp, L. N. (1993) Flood characteristics of

the Hattah Lakes system. Integrated Watering Strategy Report

No. 5, Floodplain Ecology Group, Department of Conservation

and Natural Resources, Shepparton (unpub.).

both a river and its floodplain (Jacobs 1990; Boulton

& Lloyd 1992). Environmental flows may be used to

restore wetlands which have suffered reduced

flooding as a result of regulation (Petts 1996; Tharme

2003). The RAMSAR listed Hattah Lakes are a

system of 17 floodplain lakes on the mid to lower

River Murray targeted to receive environmental flows

for ecological gain (Siebentritt'!). Regulation has

caused a decrease in the flooding frequency of one of

the first lakes in the system, Lake Hattah (Barker?;

Robinson 1966; Shaw?; Cumming & Lloyd?) whilst

the effect of regulation on the other 16 lakes is

unknown. If the maximum environmental benefit is

to be gained by providing environmental flows to the

Hattah Lakes, an estimate of the impact of regulation

on the flooding frequency of the other lakes within

the system is required. This is important because the

dynamics of floodplain flooding are complex, as

many floodplain systems have intricate patterns of

connectivity and flow (Kingsford 2000). Differences

in floodplain topography can also have dramatic

effects, as small fluctuations may be the difference

between large areas of floodplain, individual lakes

and wetlands becoming inundated or remaining dry

(Taylor et al. 1996).

A previous attempt at increasing the rate of inflow

to and flooding frequency of, the lakes was made in

1972/73 when Chalka Creek, which feeds the lakes

from the River Murray, was channelised (Baker?).

The impact of this modification has not previously

been assessed.

This paper describes a novel method, using

historical records, to estimate flooding frequency

through the Hattah Lakes system. Changes in

flooding frequency of the Hattah Lakes caused by

river regulation and the modifications to Chalka

FLOOD REGIME CHANGE IN THE HATTAH LAKES VICTORIA 75

142.255°

142.32°

142.5057

Chalka

Litttle Lake

Hattahe

Victoria

Melbourne:

Hattah

Lake

-34,785%s, , Kramen ty 734.785"

wo N Oo

a ° 6

q y g

Fig. |. The Hattah Lakes system. The shaded lakes are those referred to in the text.

6 .

yy ny

0 U U i I

1908 1916 1923 1931 1939 1946 1954 1962 1969 1977 1985 1992

Year

Fig. 2. Monthly records of the height of Lake Hattah from May 1908 to October1996.

76 N. J. SOUTER

Creek are estimated and assessed. From this,

recommendations regarding future environmental

flows are suggested.

Methods

Site Description

Hattah Lakes are a series of 17 interconnected

lakes on the Victorian floodplain of the River

Murray. The system relies on overbank flow from the

Murray and is fed via Chalka Creek, an anabranch of

the river. Chalka Creek fills the northern lakes in

succession, before rejoining the Murray to the north

(Fig. 1). The lakes on the south-easterly terminal

branch fill successively at similar times to those on

the northern branch. Lake Kramen is filled via

overbank flow directly from the River. The height of

Lake Hattah, one of the first lakes in the SE terminal

branch, is directly related to the height of the River

Murray at Euston (Robinson 1966). Most lakes

dry within 12 months after the cessation of

flow (Cumming and Lloyd*). Lakes Hattah and

Mournpoul are the most permanent of the system,

holding water for several years.

To increase the rate of inflow to the Lakes, Chalka

Creek was deepened and widened in 1972/73

(Baker?). This operation was undertaken to allow

water to flow to the Hattah Lakes when discharge in

the River Murray at Euston exceeded approximately

36 700 ML/day instead of the previous approximate

level of 48 000 ML/day (Baker’).

Estimate of flooding frequency

The flooding frequencies of six of the seventeen

lakes in the system: Lakes Hattah, Mournpoul,

Konardin, Nip Nip, Bitterang and Kramen were

calculated. These lakes were chosen for their

geographical spread and because _ historical

information detailing their flooding was available

(Fig. 1).

The height, depth in metres, of Lake Hattah was

recorded monthly on a gauge from May 1908 until

April 1984 (Fig. 2). From May 1984 until October

1996 data were collected inconsistently. However

during this period the maximum height of each flood

was recorded (Hattah-Kulkyne National Park,

unpublished data). Historical information for twelve

floods which described or depicted the extent of

flooding through the system was summarised (Table

1). From 1908 until 1996 when flooding occurred it

took place in the spring/summer of each year. This

enabled the maximum Lake Hattah gauge height of

each yearly flood to be determined.

Flooding frequencies of the six lakes were

estimated by correlating the height of Lake Hattah

with the extent of flooding through the system. For

Lake Hattah to be considered to have flooded, rather

than just having received a minor inflow of water, an

arbitrary increase in lake height of 0.35 m must have

occurred, regardless of the initial lake depth. The

flood threshold for the other five lakes was estimated

as the lowest recorded Lake Hattah gauge height at

which each lake was flooded. Given this value the

number of times each lake flooded over a given

period was estimated by referring to the height of

Lake Hattah each year and determining the extent of

flooding through the system. The flooding frequency

of each lake was calculated by dividing the number

of times a lake was estimated to have flooded by the

total number of years analysed. The frequency of dry

years is equal to one minus the flooding frequency.

The completion of the Hume Dam in 1936 is

regarded in this study as the beginning of river

regulation. This provided 28 years (1908-1935) of

pre-regulation data and 61 years (1936-1996) of

post-regulation data. Two by two contingency tables

of the frequency of inflow years vs. no inflow years

for each lake were constructed to assess the impact of

regulation and Chalka Creek channelisation using a

G-test (Sokal & Rolf 1995). Three comparisons

between flooding frequency for each lake were

made:

1) Pre-regulation (1908-1935: 28 years) vs. the

period after regulation but prior to the

channelisation of Chalka Creek (1936-1972:

37 years),

2) Post-regulation before (1936-1972) vs. after the

channelisation of Chalka Creek (1973-1996:

24 years), and

3) Pre-regulation (1908-1935) vs. post regulation

after the channelisation of Chalka Creek (1973-

1996).

Results

Flooding characteristics

The determination of flood threshold values for

each of the six lakes are:

LAKE HATTAH

The lack of records describing floods which

exclusively filled Lake Hattah made it difficult to

determine what level of inflow to Lake Hattah

constitutes a flood or a minor inflow of water. On

only four occasions since 1908 did water enter Lake

Hattah but not fill to a level likely to move further

through the system. The lake rose 0.10 m, (from 2.28

to 2.38 m) in 1910, 0.30 m (from 1.98 to 2.28 m) in

1922, 1.78 m (from 0.51 to 2.29 m) in 1928, whilst

the 1987 flood entered the lake but its height did not

increase (Table 1). In calculating flooding frequency

only 1928 was analysed as a flood, whilst the minor

inflows in 1910, 1922 and 1987 were classified as

years of no inflow.

FLOOD REGIME CHANGE IN THE HATTAH LAKES VICTORIA 77

TABLE 1. Summarised historical records of flooding through the Hattah Lakes system, showing the terminal lake flooded

along each arm of the system and the maximum height of Lake Hattah for each flood.

Source Year Terminal Lake = Terminal Lake = Maximum height of

flooded (northern arm) (eastern arm) Lake Hattah

Department of Natural Resources 1956 Bitterang Kramen 5.49 m

and Environment (DNRE)>

Robinson (1966) 1964 Bitterang Nip Nip 4.50 m

Robinson (1966)° 1964 Bitterang 40m

Robinson (1966)° 1964 Nip Nip 3.66 m

Douglas’ 1970 Bitterang Not reported 4.27 m

DNRE® 1986 Mournpoul Hattah 2.74 m

DNRE® 1987 Lockie Hattah inflow but no height increase

DNRE® 1989 Bitterang Nip Nip 3.75 m

TM Satellite photograph (15 Dec 1989) 1989 Bitterang Nip Nip 3.75 m

DNRE?® 1990 Bitterang N/R 3.76 m

MDBC (1994)!° 1990 Bitterang Nip Nip 3.76m

TM Satellite photograph (9 Feb 1993) 1992 Bitterang Nip Nip 4.59m

(minimal input

to Kramen)

DNRE? 1993 Bitterang Kramen 5.0m

DNRE'"! 1995 Konardin Nip Nip 3.60 m

(reached but

did not fill)

Puckridge ef al.'* September Mournpoul - <2.57m (actively flooding

1996 when recorded)

Puckridge ef al.'? October Bitterang Nip Nip 3.95 m

1996

5 Aerial photograph (unpub.)

° Robinson reported that Lake Bitterang began to fill once the height of Lake Hattah reached 4.0 m, whilst for

Lake Nip Nip to fill Lake Hattah must reach 3.66 m.

7 Douglas, M. G. (1972) The water regime of the Chalka Creek and lakes of the Kulkyne Forest and Hattah Lakes

National Park. Sunraysia Naturalists’ Research Trust - Ninth Report (unpub.).

& Compiled during the preparation of Cumming and Lloyd*

° Brendan Atkins (pers. comm.)

'0 Terrascan Aerial photograph

'! Phil Murdoch (pers. comm.)

Puckridge, J. T., Ward, K. A. and Walker, K. F. (1997) Hydrological determinants of fish and macroinvertebrate ecology

in the Hattah Lakes System: Implications for Time-Share Flooding. Part 1: 1996/97. University of Adelaide,

Department of Natural Resources and Environment, Land and Water Resources Research and Development

Corporation (unpub.).

LAKE MOURNPOUL

Lake Mournpoul was found to require a slightly

larger flood than Lake Hattah for it to fill. Water was

observed to reach Lake Hattah before Lake

Mournpoul in 1970, 1986 and 1989, whilst in 1987

water did not enter Lake Mournpoul but flowed into

Lake Hattah. The 1986 flood reached a gauge height

of 2.74 m and filled only Lakes Hattah and

Mournpoul. Puckridge ef. al/.'* noted that in

September 1996 Lake Mournpoul was actively

filling when Lake Hattah had a gauge height of 2.57

m and that Lake Mournpoul had probably flooded

earlier as a result of an earlier unobserved flood

peak. Using this information the flood threshold of

Lake Mournpoul has been estimated at 2.57 m on the

Lake Hattah gauge.

LAKE KONARDIN

In 1995 Lake Konardin flooded when Lake Hattah

was at a height of 3.60 m and was the last lake

flooded on the northern arm of the system. Prior to

flooding the Lake was dry (Phil Murdoch, Hattah-

Kulkyne National Park Ranger pers. com.). The

flood threshold of Lake Kramen is estimated at 3.60

m on the Lake Hattah gauge.

LAKE Nip Nip

Robinson (1966) determined that Lake Hattah

must reach about 12’ (3.66 m) before Lake Nip Nip

floods. This observation was supported in 1995

when Lake Hattah rose to 3.60 m and water reached,

but did not fill Lake Nip Nip. The floods of 1989,

1990, 1992 and 1996 all showed lake Nip Nip to fill

78 N. J. SOUTER

TABLE 2.

Lake Hattah gauge height and frequencies of no inflow and flooding for six lakes in the Hattah system.

Frequencies are presented for a range of time spans: pre-regulation and post regulation both before and after Chalka

Creek channelisation.

No Inflow Hattah Mournpoul Konardin Nip Nip Bitterang Kramen

Lake Hattah gauge height (m) - - 2.75 3.60 3.66 3.75 5.0

Pre-regulation (1908-1935) 0.32 0.68 0.64 0.50 0.50 0.43 0.07

frequency

Post-regulation/pre-Chalka 0.59 0.41 0.41 0.32 0.32 0.32 0.03

modification (1936-1972)

frequency

Post-regulation/post-Chalka 0.33 0.67 0.67 0.38 0.33 0.33 0.13

modification (1973-1996)

frequency

TABLE 3. G-test results for comparisons of the flood frequency of the Hattah lakes for periods pre- (1908-1935), and post-

regulation prior to (1936-1972) and after (1973-1996) Chalka Creek channelisation (* significant difference at the

p<0.05 level).

Lake 1908-1935 vs. 1936-1972 1936-1972 vs. 1973-1996 1908-1935 vs. 1973-1996

= p= o p= is p=

Hattah 4.733 0.030* 3.931 0.047* 0.008 0.928

Mournpoul 3.550 0.059 3.931 0.047* 0.031 0.859

Konardin 1.999 0.157 0.160 0.689 0.798 0.371

Nip Nip 1.999 0.157 0.005 0.942 1.439 0.230

Bitterang 0.722 0.395 0.005 0.942 0.482 0.487

Kramen 0.606 0.436 1.984 0.159 0.387 0.534

when the gauge height of Lake Hattah was over 3.66

m (Table 1). The threshold of Lake Nip Nip is

estimated at 3.66 m on the Lake Hattah gauge.

LAKE BITTERANG

Robinson (1966) reported that Lake Bitterang

began to fill when Lake Hattah reached 4.0 m. This

is considerably higher than the values recorded

during 1989, 1990 and 1996. This difference cannot

be explained by Lake Hattah holding water

previously, as prior to the 1964 flood it had been dry

for approximately ten months. The flood threshold of

Lake Bitterang 1s likely to be no greater than the 3.75

m recorded for the 1989 flood and verified via two

sources (Table 1). However, the threshold must be

greater than 3.60 m, the height of the 1995 flood,

which did not reach Lake Bitterang, suggesting that

the true threshold value lies somewhere between 3.60

and 3.75 m. The threshold value has been estimated

as 3.75 m on the Lake Hattah gauge. The other four

floods to fill Lake Bitterang (1956, 1970, 1992 and

1993) were all well above this value.

LAKE KRAMEN

The threshold value for Lake Kramen lies

somewhere between 4.59 and 5.0 m as the 1992

flood of 4.59 m reached Lake Kramen but did not fill

it, whilst the 1993 flood of 5.0 m filled the Lake. The

aerial photograph of the 1956 flood revealed

significant connection directly between the River

Murray and Lake Kramen, suggesting that the Lake,

when it fills, receives most of its water directly from

the River Murray and not via the Chalka Creek

system. The threshold of Lake Kramen is estimated

at 5.0 m. Whilst there was some difference between

the two values used to estimate this threshold, the use

of the higher value is justified in that on only two

occasions (1939, 4.65 m and 1973, 4.88 m) did

floods of a height between these values occur.

Changes in flooding frequency

Prior to regulation of the River Murray, the Hattah

Lakes system was subject to a wide range of flooding

frequencies (Table 2). Lakes Hattah and Mourpoul

were the most frequently flooded, Lakes Konardin,

Bitterang and Nip Nip formed a group of

intermittently flooded lakes and Lake Kramen was

only infrequently filled. Post-regulation (1936-1972)

flooding frequencies for all lakes were lower than

those estimated prior to regulation (Table 2). The

flooding frequency of Lake Hattah was significantly

lower, whilst the frequencies of other lakes were not

significantly different (Table 3). Post-regulation the

flooding frequencies of lakes Hattah and Mournpoul

FLOOD REGIME CHANGE IN THE HATTAH LAKES VICTORIA 79

were identical, as were the frequencies of lakes

Konardin, Bitterang and Nip Nip. There was little

difference in the flooding frequencies of Lake

Kramen either before or after regulation

Lakes Hattah and Mournpoul were flooded

significantly more frequently after the channelisation

of Chalka Creek than in the 37 years prior (Table 3).

Channelisation significantly increased the flooding

frequency of Lake Hattah which returned to its pre-

regulation level, as there was no. significant

difference in the flooding frequency prior to

regulation and post Chalka Creek modification

(Table 3). Channelisation did not increase flooding

frequency within the rest of the system as there was

little change in the flooding frequencies of Lakes

Konardin, Nip Nip and Bitterang, nor were

differences in flooding frequency between lakes

significant (Table 3).

Discussion

Regulation of the River Murray has reduced the

frequency and variability of flooding within the

Hattah Lakes. This was primarily due to the

reduction in the frequency of small to medium sized

floods. This in turn reduced variability as lakes

which could be filled by small to medium sized

floods were now only generally filled via larger

floods which filled all of the lakes. This investigation

arrived at the same conclusion as the River Murray

monthly simulation model of Maheshwari ef al.

(1995) which showed that regulation had caused a

reduction in the frequency of small to medium sized

floods in the mid-Murray. Given the positive

relationship between the height of Lake Hattah and

the River Murray at Euston (Robinson 1966), this

result was not unexpected. Kingsford (2000)

suggested that instream models are unsuitable for

assessing flood regime on a river’s floodplain. The

value of investigating the effect of river regulation on

the floodplain, rather than just the main channel, was

demonstrated by the findings of this study that

regulation reduced the variability of flooding

frequencies between lakes, a result of the reduction

of small to medium sized floods.

The channelisation of Chalka Creek in 1972/73

did, as predicted (Baker?; Shaw?), increase the

flooding frequencies of lakes Hattah and Mournpoul

(even restoring the pre-regulation flooding frequency

of Lake Hattah). Channelisation had no impact on

the systems less frequently flooded lakes and did not

fully restore the variability of flooding through the

system.

Whilst a loss of variability in the flood regime was

observed, differences in flooding frequency between

periods for any of the lakes were generally not

statistically significant. This does not necessarily

mean that non-statistically significant changes may

not be biologically important as large differences in

flooding were required for significance. The

relationship between flooding and the biological

response in the Hattah Lakes is beyond this paper

and will be addressed elsewhere. The inability to

detect statistical differences in the flooding

frequencies between lakes could also be due to the

short time intervals examined. The longest period of

37 years (between 1936-1972) saw a considerable

increase in the degree of regulation of the River

(Maheshwari et al. 1995) added to which the flow

regime of the River Murray is unstable within a 50

year time frame (Walker ef al. 1995). Thus it is likely

that larger data sets, although unavailable, would be

required to achieve significant results.

Caution in these results should also be exercised, as

sources of error exist in determining the flood

threshold height for each lake. It is likely that

multiple flood threshold values exist for each of the

lakes. For example, threshold values may change

according to the speed at which flooding progresses

through the system. As this information was

unavailable or inadequate it could not be assessed.

Differences in the degree of drawdown at the start of

each flood may also alter the threshold height.

However most of the lakes within the system are

likely to have dried between floods as most are

shallow and dry within twelve months after the

cessation of flow (Cumming and Lloyd*). Lakes

Mournpoul (drying time, 7 years) and Hattah (3

years) are two of the deeper lakes in the system and,

taking longer to dry (Cumming and Lloyd?) often

have different drawdown conditions at the start of

each flood. These conditions were assessed in

determining the flooding frequency of Lake Hattah

although they were unknown for Lake Mournpoul.

Further detailed on ground investigations of flooding

through the system are recommended to improve the

accuracy of the flooding frequency estimates

provided in this paper.

The loss of flooding variability may have

significant ecological impacts, given the importance

of flooding frequency in structuring aquatic

communities. With the error associated in

calculating flooding frequency these values should

be used as a guide only. However, it can be

reasonably suggested that efforts to rehabilitate the

Hattah Lakes, through the provision of

environmental flows, would be best directed towards

increasing the variability of flooding through the

system, most particularly within intermittently

flooded lakes such as Konardin, Nip Nip and

Bitterang as these lakes have suffered reduced

flooding frequency and variability as a result of

regulation but have not been improved by the

modifications to Chalka Creek.

80 N. J. SOUTER

Acknowledgements

This work arose out of the Land and Water

Resources Research & Development Corporation,

project VCB1, Time-share flooding of aquatic

ecosystems. Keith Ward, Jim Puckridge, Alex

McNee and Lance Lloyd are thanked for their

support and advice on earlier drafts. Trevor

Jacobs suggested that I prepare this work for

publication.

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BRIEF COMMUNICATION

HELMINTH AND PROTOZOAN PARASITES OF FERAL CATS

FROM KANGAROO ISLAND

There are limited data on the helminth and protozoan

parasites of the feral cat, Felis catus, on Kangaroo Island

although the carnivore is well established there. Five

helminth species from feral cats were reported in a list

compiled from parasites that had been submitted to a

veterinary laboratory in South Australia'. Surveys of the

helminth species in feral cats conducted elsewhere have

identified a varying prevalence dependent on the available

food sources and climate*+5°7, Few of these surveys,

however, identified the protozoan parasites. A significantly

greater prevalence of the protozoan parasite, Toxoplasma

gondii, has been identified in sheep on Kangaroo Island and

attributed to an apparent presence of large numbers of feral

cats*. In this study, we present data on the occurrence of the

helminth and protozoan parasites detected in a sample of

feral cats from Kangaroo Island.

The gastro-intestinal tract, heart, lungs, whole blood and

faeces from 46 cats were submitted for examination.

Organs were opened, washed and the parasites removed and

counted using a dissecting microscope. The stomach wall

was examined for nodules. Mucus from the stomach wall

was examined specifically for Ollulanus tricuspis using a

squash preparation on a glass slide and a compound

microscope. The stomach and small intestinal wall was

scraped, fixed in formalin and examined for helminths. A

faecal sample was examined for helminth eggs, nematode

larvae and protozoan cysts using a centrifugal flotation

method in saturated MgSO, solution and additionally in

saturated KI solution to determine the presence of

Cryptosporidium oocysts’. The Baermann method! was

used to recover Aelurostrongylus abstrusus larvae from

faeces and lung parenchyma. Faecal samples were stored in

2% K2Cr,O; so that coccidia could sporulate for

identification. Nematodes were preserved in alcohol,

cleared in lactophenol and identified using a compound

microscope. Armed scoleces of preserved Tuenia

specimens were removed, mounted and cleared in

DeFauré’s medium for the examination of rostellar hooks.

Identification of Zaenia specimens was made by counting

and measuring the large and small rostellar hooks and

TABLE 1. Helminth and protozoan parasites found in 46 feral cats from Kangaroo Island.

Parasite Site Prevalence (%) Abundance (mean)

Acanthocephala

Oncicola pomatostomi Intestine 7 2-7 (4)

Nematoda

Aelurostrongylus abstrusus Lung 11 -

Ancylostoma tubaeforme Intestine 15 1-16 (4)

Cyathospirura dasyuridis Stomach 15 1-78 (98)

Cylicospirura felineus Stomach 57 1-39 (7)

Ollulanus tricuspis Stomach 2 6

Toxocara cati Intestine 76 1-53 (9)

Cestoda

Dipylidium caninum Intestine 4 25-31 (28)

Spirometra erinacei Intestine 39 1-26 (5)

Taenia taeniaeformis Intestine 63 1-31 (5)

Trematoda

Brachylaima cribbi Intestine 2 1

Protozoa Faeces

Cryptosporidium f4

Giardia 2

Isospora felis 15

Isospora rivolta 4

Sarcocystis 7

Toxoplasma (serology) IHAT 87

TABLE 2. Comparison between the detection of helminths in the intestine and the detection of eggs in faecal samples.

Helminth Intestine Faeces

Number infected Positive for eggs

Ancylostoma tubaeforme 7 3

Brachylaima cribbi ] 1

Cyathospirura dasyuridis &

Cylicospirura felineus 26 4

Oncicola pomatostomi 3 0

Spirometra erinacei 18 14

Toxocara cati 35 28

Taenia taeniaeformis 29 13

comparing the data with those of Verster'!. An indirect the prevalence of these two parasites may be

haemagglutination test (Toxo HAI commercial test kit

{Fumouze]) and a direct agglutination test (Antigene Toxo-

AD commercial test kit [bioMéricux]) were used for

detecting Toxoplasma antibody in serum samples from 47

cats. A dilution of 1:80 was regarded as positive for the HAI

test and 1:4 for the DA test following the manufacturers

instructions.

All of the cats examined were mature; six were regarded

as young adults. Twenty-one cats were female and 25 male.

Ten of the female cats were pregnant and two were in

lactation.

The majority of the parasites found (Table 1) have been

reported previously in surveys of feral cats. This study

confirms that Tuenia taeniaeformis, Spirometra erinacei

and Toxocara cati are common parasites of feral cats.

Ancylostoma tubaeforme is regarded as the common

hookworm of cats and has been reported in feral cats from

the Northern Territory’ and from around Sydney’.

Cylicospirura felineus and Cyathospirura dasyuridis were

found in tumour-like nodules on the stomach wall.

Cylicospirura felineus was the predominant species and

occurred together with Cyathospirura dasyuridis in seven

cats. The trematode, Brachylaima cribbi, occurs in a variety

of mammals and birds, has helicid snails as intermediate

hosts and is infectious to man!?. Oncicola pomatostomi was

detected in 65% of 188 feral cats from the Northern

Territory suggesting that birds, as paratenic hosts,

constituted a significant part of their diet’. In this study, the

parasite was less common, however there is no information

on the abundance of this parasite in birds from Kangaroo

Island preventing further inference. The prevalence of

Isospora felis was higher than that reported in feral cats

elsewhere. Infections by /sospora spp. are considered age

dependent, occurring more commonly in younger

animals*5, Whilst Giardia sp. was found in a faecal sample

from only one cat in this study, higher prevalence reported

elsewhere! has implicated the feral cat in the transmission

of the parasite to wildlife and man. It has been suggested

that Cryptosporidium sp. recovered from domestic cats is

not of zoonotic significance and the oocysts are smaller

than those recovered from man'*. Furthermore, cats appear

to carry different species such as C. baileyi and C. muris'*

essentially from birds and rodents respectively. We were

unable to determine the identity of the species detected in

this study. McGlade ef a/.'> reported a significantly greater

prevalence of Cryptosporidium sp, and Giardia sp. in cats

using PCR in comparison to microscopy and consequently

underestimated here. The dimensions of Sarcocystis sp.

sporocysts here are consistent with those of S. gigantea

(x length 13.2 tm x ¥ width 9.4 um, n = 22), a species

infecting sheep. Faecal examination may also underestimate

the prevalence of toxoplasmosis. The shedding of

Toxoplasma gondii oocysts in faeces occurs for a short time

and only once following infection, usually when the young

cat begins hunting rodents and birds'*. Serological tests

indicate previous exposure to the parasite. The high

prevalence of Toxoplasma antibodies in feral cats is

consistent with the high prevalence of antibodies in sheep

on Kangaroo Island.

There was a poor correlation between the detection of

adult nematodes and the detection of nematode eggs in

faecal samples (Table 2). Zoxocara cati eggs were not

detected in faecal samples on seven occasions, six being

due to infections with immature nematodes. Hookworm

eggs were detected in the faeces of three of the seven cats

infected. Of the four negative samples, three contained a

single worm and the fourth contained three, non-gravid

female worms. The detection of cestode and

acanthocephalan eggs in faeces of cats infected with adult

worms was similarly inconsistent. Infections of up to 31

cestodes remained undetected by faecal examination. These

results indicate that coprological surveys for helminth

parasites may underestimate the prevalence of infections.

Aelurostrongylus abstrusus are small worms, <10 mm long,

occurring in lung parenchyma. Adults were not recovered

from lungs when examined macroscopically, however,

using the Baermann technique, Aelurostrongylus larvae

were recovered from lung tissue and faecal samples of

positive cats.

The information presented in this study identified similar

helminth and protozoan parasites in feral cats from

Kangaroo Island to those detected elsewhere in Australia

and confirms that feral cats may act as reservoirs for these

parasites. The study suggests that the feral cat is responsible

for the high prevalence of Yoxoplasma in sheep on

Kangaroo Island and poses a potential disease risk to

wildlife. It also establishes that feral cats in Australia are

infected with Cryptosoporidium.

'O’Callaghan, M. G., Moore, E. & Ford, G. E. (1984) Helminth

and arthropod parasites from dogs and cats in South Australia. Aust.

vet. Practit. 14, 159-161.

2Coman, B. J. (1972) A survey of the gastro-intestinal parasites of

the feral cat in Victoria. Aust. vet. J. 48, 133-136.

‘Ryan, G. E. (1976) Gastro-intestinal parasites of feral cats in New

South Wales. Aust. vet. J. 52, 224-227.

‘Gregory, G.G. & Munday, B. L. (1976) Internal parasites of feral

cats from the Tasmanian midlands and King Island. Aust. vet. J. 52,

317-320.

°Coman, B. J., Jones, E. H. & Driesen, M. A. (1981) Helminth

parasites and arthropods of feral cats. Aust. vet. J. 57, 324-327.

°Coman, B. J., Jones, E. H. & Westbury, H. A. (1981) Protozoan

and viral infections of feral cats. Aust. vet. J. 57, 319-323.

'O’Callaghan, M. G. & Beveridge, I. (1996) Gastro-intestinal

parasites of feral cats in the Northern Territory. Trans. R. Soc. S.

Aust. 129, 175-176.

‘O’Donoghue, P. J., Riley, M. J. & Clarke, J. F. (1987) Serological

survey for 7oxoplasma infections in sheep. Aust. vet. J. 64, 40-45.

°O’ Donoghue, P. J. (1995) Cryptosporidium and cryptosporidiosis

in man and animals. Int. J. Parasitol. 25, 139-195.

"Georgi, J. R. (1974) Parasitology for Veterinarians (W. B.

Saunders, Philadelphia. USA.) pp 134-135.

"Verster, A. (1969) A taxonomic revision of the genus Taenia

Linnaeus, 1758 s. str, Onderstepoort J. vet. Res. 36, 3-58.

"Butcher, A. R. & Grove, D. I. (2001) Description of the life-cycle

stages of Brachylaima cribbi n. sp. (Digenea: Brachylaimidae)

derived from eggs recovered from human faeces in Australia. Sys.

Parasitol. 49, 211-221.

Milstein, T. C. & Goldsmith, J. M. (1997) Parasites of feral cats

from southern Tasmania and their potential significance. Aust. ver. J.

75, 218-219.

“Sargent, K. D., Morgan, U. M., Elliot, A. & Thompson, R. C. A.

(1998) Morphological and genetic characterisation of

Cryptosporidium oocysts from domestic cats. Vet. Parasitol. 77,

221-227.

'McGlade, T. R., Robertson, I. D. Elliot, A. D., Read, C. &

Thompson, R. C. A. (2003) Gastro-intestinal parasites of domestic

cats in Perth, Western Australia. Vet. Parasitol. 117, 251-262.

‘Dubey, J. P. & Beattie, C. P. (1988) Toxoplasmosis of animals and

man. (CRC Press Boca Raton, Florida. USA) pp 18-19.

M. O’CALLAGHAN, South Australian Research and Development Institute, GPO Box 397, Adelaide, SA

5001, E-mail: ocallaghan.micko@saugov.sa.gov.au, J. REDDIN, Primary Industries and Resources, PO Box

469 Murray Bridge, SA 5253 and D. LEHMANN, Kangaroo Island Veterinary Clinic, Kingscote, SA 5223.

eo 6)VOL. 129, PART 2

30 NOVEMBER, 2005

RSA .

A special issue of the Transactions of the

Royal Society of South

Australia Incorporated

containing papers on the 2004 Expedition

to the Althorpe Islands, South Australia

Guest Editors: Sue Murray-Jones (BSc, PhD)

Scoresby Shepherd (B.A., LL.B., M.Env.St., Ph.D.)

INCORPORATING THE

Records of the

South Australian Museum

Contents.

Murray-Jones, S. & Shepherd, S. A. An expedition to the Althorpe Islands, South

Australia: Introductory narrative and conservation

recommendations

Zang, W. L. Geology of Althorpe Island

Radford, A. Human settlement on Althorpe Island and condition of the

lighthouse complex -

Lawley, E. F. & Shepherd, S. A. Land use and vegetation of Althorpe Island, South

Australia, and a floristic comparison with South Neptune Islands

Lawley, E. F., Lawley, J. J. & Page, B. Effects of African boxthorn removal on native

vegetation and burrowing of short-tailed shearwaters on Althorpe

Island, South Australia _-

Baldock, R. N, & Womersley, H. B. S. Marine benthic algae of the Althorpe Islands,

South Australia

Baker, J. L., Edgar, G. J. & Barrett, N.S. Subtidal macroflora of Althorpe and Haystack

Islands, South Australia

Benkendorff, K. Intertidal molluscan and echinoderm diversity at Althorpe Islands

and Innes National Park, South Australia

Staples, D. A. Pycnogonida of the Althorpe Islands, South Australia -— —

Walker-Smith, G. K. A new species of Neopeltopsis (Copepoda, Harpacticoida,

Peltidiidae) from Althorpe Island, South Australia -— — -

Shepherd, S. A., Edgar, G. J. & Barrett, N. S. Reef fishes of the Althorpe Islands and

adjacent coasts of central South Australia

Shepherd, S. A., Teale, J. & Muirhead, D. Cleaning symbiosis among inshore fishes at

Althorpe Island, South Australia and elsewhere -— - -— —

Shepherd, S.A. & Brook, J. B. Foraging ecology of the western blue eases Achoerodus

gouldii, at the Althorpe Islands, South Australia — -

Einoder, L. D. & Goldsworthy, S. D. Foraging flights of short-tailed shearwaters

(Puffinus tenuirostris) from Althorpe Island: assessing their use of

neritic waters

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

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Transactions of the Royal Society of S. Aust. (2005), 129(2), 85-89.

AN EXPEDITION TO THE ALTHORPE ISLANDS, SOUTH AUSTRALIA:

INTRODUCTORY NARRATIVE AND CONSERVATION RECOMMENDATIONS

by S. MurrAy-JONES* & S. A. SHEPHERD*

Summary

Murray-Jones, S, & SHEPHERD, S. A. (2005) An expedition to the Althorpe Islands, South Australia:

Introductory narrative and conservation recommendations. Trans. R. Soc. S. Aust. 129(2), 85-89, 30 November,

2005.

A general overview of the 2004 Expedition to the Althorpe Islands group is provided, including a description

of the islands, an outline of the research program, a brief introduction to the papers presented in this volume,

and some recommendations for management.

Key Worps: Southern Australia, Althorpe Is, lighthouse, geology, terrestrial ecology, heritage, marine

conservation.

Description of islands and habitats

The Althorpe Is consist of one main island and five

small islets (Fig. 1), of which Althorpe I. is the

southernmost and largest (96 ha, and 93m high). The

five small islets (the Western Isles) are off the

western side of Althorpe I. (Baker 2005).

To the NE, and closer to the coast, are Seal |., and

Haystack I. (43 m high). Together, the islands and

islets are informally known as the Althorpe Is Group.

The islands are the remains of once prominent

granite hills surmounted by Bridgewater calcarenite.

The islands were probably isolated from the

mainland some 9 — 10 000 yrs ago (Zang 2005).

All islands in the Althorpe group have sub-tidal

granite basement reefs, with some calcarenite blocks

present in the nearshore area in some places, where

they have eroded from the island “capping” above.

On Althorpe I., jointing in the massive granite humps

has resulted in gaping, many-branched crevasses and

chasms along the coastline, that alternate with long

fingers of rock, in turn dissected into segmented

chains of islets. The small Western Isles are the

eroded and segmented remnants of a granite rise

along a series of joints or dykes (Robinson ef al.

1996). Mooring Bay, bounded by cliffs on the NE

side of Althorpe I., provides the only anchorage

* Coast Protection Branch, Natural & Cultural Heritage,

Department for Environment & Heritage (DEH), GPO Box 1047,

Adelaide, South Australia 5001.

* South Australian Research and Development Institute (SARDI),

Aquatic Sciences, PO Box 120, Henley Beach, South Australia

$022.

' The names given in Fig. | were gazetted in the SA Government

Gazette on 13th Oct. 2005 (p3685). The main island of the

Althorpe Is group (commonly known as Althorpe I.) is in fact

unnamed. A proposal to name it Laubadere I., after the name

given to it by Freycinet in 1824, is under consideration by the

Geographical Names Advisory Committee of the Government of

SA.

(Robinson ef al. 1996). The names adopted in Fig. |

are mainly those in use by lighthouse staff and the

Friends of the Althorpe Is Conservation Park, with a

few adopted by the 2004 expedition. All names have

been accepted by the Geographical Names Advisory

Committee and await ratification!.

Althorpe Islands

The base of the main island is made up of granite

rock rising up to ~15 m above sea level. Sitting on

top of this granite base are layers of sandstone, which

are the remains of ancient sand dune systems that

once covered the region. All this is capped by layers

of calcarenite. The geology of Althorpe I. is

described in more detail in Wang (2005, this

volume).

Althorpe I. rises steeply to a plateau 93 m above

sea level. On the plateau there is an abandoned

airstrip, a lighthouse, and three lighthouse keepers’

cottages. The northern islet of the Western Isles

actually consists of four islets separated by narrow

channels. The base of all four islets is granitic, with

the two middle islets capped with calcarenite. These

form two peaks rising to 23 m and 22 m respectively

above sea level. The two outer islets are bare granite

only. The southern islet, also informally called ‘Tern

I” consists of bare granite projecting 12 m above sea

level.

Haystack Island

This island has a narrow wall of sheer cliffs, made

of sandstone that has been undermined and indented.

The cliffs sit on a supporting ridge of granite that lies

well below sea level. The island has been eroded into

a series of tall lobes connected by thin necks of rock,

two of the lobes rising to 44 m and 41 m respectively

above sea level. The sandstone is capped with

limestone, which forms a series of stepped terraces

on top of the domes (Robinson e¢ al. 1996).

86 S. MURRAY-JONES & S, A. SHEPHERD

Stenhouse Bay

Innes NP :

Haystack Island

Seal Island

__Althorpe Island

Fig. 1. Map of the Althorpe Islands Group.

Seal Island

This island rises 35 m above sea level on a hump

of granitic gneiss, intruded by amphibolite dykes.

The sea has eroded deep grooves and indentations

that channel the swell into a surge that reaches the

island platform of sandstone capped with

limestone. These surges undermine the limestone

cap, the fringe of which is strewn with jagged

fragments of rock from collapsed overhangs. The

broadest joints and dykes have been worn to deep

indentations, or completely penetrated, to form the

segmented islets off the western tip of Seal I.

(Robinson et al. 1996).

Marine habitat

The habitat diversity in the area includes: near-

shore benthic granitic basement reefs with a diversity

of forms such as platforms, ledges, boulders, caves,

chasms and crevasses, overhangs, vertical rock walls,

isolated reef outcrops and near-shore fringing reef,

patches of broken calcarenite blocks and rubble;

areas of small sandy beaches in Mooring Bay; mixed

sand / granite reef / calcarenite block / rubble reef

intertidal habitats; subtidal seagrass beds (e.g. NE

Althorpe I. and Haystack I.); and benthic sand

habitat (Robinson et al. 1996; Edyvane 1999;

Edyvane & Baker 1998).

2004 Expedition

The Royal Society of South Australia, with a long

history of sponsoring expeditions to the State’s

offshore islands (Robinson ef al. 2003), promoted

this expedition, with major funding support by South

Australian Research and Development Institute and

Department for Environment and Heritage. Previous

visits to the Althorpe Is include some bird

observations by Perryman (1937), and a brief survey

by a National Parks & Wildlife SA party on 24 — 25

Nov. 1982 of the plants, mammals, birds and reptiles

(Robinson et al. 1996), Since 1982, the Friends of

the Althorpe Is Conservation Park (FoATCP) have

INTRODUCTORY NARRATIVE & CONSERVATION RECOMMENDATIONS ON ALTHORPE ISLANDS, SA 87

made regular visits to the island, and accumulated

much additional data (e.g. Lawley & Shepherd

2005).

The scientific program that is the focus of this

special edition took place from 29 Jan. to 12 Feb.

2004, and comprised a terrestrial and a marine

component. The terrestrial party occupied the

lighthouse cottages on Althorpe I., with members of

FoAICP, while the marine party stayed either aboard

the fisheries research vessel Ngerin, present from | —

8 Feb. 2004, or on shore. Small boats were used to

ferry scientists between islands. The following

persons took part in the expedition’s scientific

programs:

Terrestrial program

Alison Radford (land-use history and

archaeology), assisted by Vicki Cheshire; Rob

Fitzpatrick (soils) assisted by Alison Fitzpatrick;

Brad Page and Shelley Harrison (seabird feeding),

assisted by Vicki Cheshire and Erika Lawley;

vegetation surveys (Erika and John Lawley).

Marine program

Robert Baldock, (algal systematics), Anthony

Cheshire (algal ecophysiology); Bayden Russell and

Jarrod Stehbens (ecology of marine macroalgae);

Scoresby Shepherd (feeding ecology of groper;

cleaning behaviour of fishes); Kirsten Bilgmann and

Sue Gibbs (dolphin genetics and habitat use);

Alastair Hirst (macroepifauna of algae); Rob Lewis

(rock lobster population assessment, assisted by

James Brook); David Staples (systematics of

pycnogonids); Chris Halstead, Graham Edgar,

Neville Barrett, Ali Bloomfield and James Brook

(biodiversity assessment); Thierry Laperousaz

(animal collections for SA Museum); Kirsten

Benkendorff and Alex Gaut (intertidal biodiversity).

Sue Murray-Jones assumed responsibility for the

logistics and assisted in the algal ecophysiological,

fish and dolphin habitat work.

Friends of Althorpe Islands Conservation Park

(FoAICP)

FoAICP was formed in 1996, after attempts by

DEH to find a lessee for the Island failed. The

current membership is about 150. The Friends

undertake quarterly visits to the island throughout

the year to carry out maintenance on buildings, the

solar/wind power energy system, equipment and

pathways, and other tasks such as weed and feral

animal control. Funding for these is usually obtained

through project grants.

Participants and their roles included: John Lawley

(island caretaker, logistics and coxswain duties); Les

Harper (coast guard radio); John Webster and David

White (cottage cleaning and preparation, water,

electricity and mouse control); Erika Lawley

(kitchen coordination, guide to island locations,

shearwater research assistant); Stefania Madonna

and Christine Lawley (domestic duties, terrestrial

work, weed control).

Operations support

Dave Kerr (Master of Ngerin), Chris Small (Ngerin

Mate/Engineer), and Michael Clark (Dive

Supervisor), all from SARDI; and from DEH, Rick

James (acting Mate); Danny Doyle, Caroline

Paterson, and Tim Collins (general support), as well

as support from FoAICP.

Results

The terrestrial studies in this special issue have

consolidated historical and biological information

not generally accessible. These include: the history

of human occupation of Althorpe I. and the condition

of the lighthouse complex (Radford 2005); and a

brief history of the land use of the island, with its

likely effects on the vegetation (Lawley & Shepherd

2005). The response of the island’s vegetation and

the nesting of the short-tailed shearwater, Puffinus

tenuirostris following removal of an African

boxthorn infestation is considered by Lawley ef al.

(2005), while Einoder & Goldsworthy (2005)

describe the at-sea movement and marine habitat use

of the shearwaters. In addition, a description of the

geology (Zang 2005) of Althorpe I. is included,

although the study was done during independent

visits to the Island.

The marine studies in this issue have substantially

extended knowledge of benthic and intertidal

communities of the region. The diversity of

intertidal molluscs and echinoderms is described by

Benkendorff (2005). Staples (2005) describes the

pycnogonid fauna of the area, including two new

species, Pseudopallene watsonae and P. inflatus.

Walker-Smith (2005) describes a new species of

Neopeltopsis and documents the occurrence of other

species of Peltidiidae from Althorpe I. Baldock &

Womersley (2005) summarize past and recent algal

collections by providing a species list of benthic

algae. Baker et al. (2005) describe the algal diversity

and community structure at Althorpe and Haystack

Is, using data collected both in 1993 and during the

2004 expedition. A description of the reef fishes of

Althorpe and Haystack Is, as well as of some

mainland control sites, is provided by Shepherd er

al. (2005a). Shepherd & Brook (2005) describe the

feeding behaviour of Western Blue Groper, while

Shepherd ef al. (2005b) describe cleaning symbioses

amongst inshore fishes, both for the Althorpe Is and

elsewhere, the first such account for South

Australia.

88 S. MURRAY-JONES & S. A. SHEPHERD

Recommendations

After the boxthorn removal program described by

Lawley et al, (2005), native vegetation was found to

recolonise bare patches, and there is some evidence

that shearwater burrows may have expanded into

bare areas as well. Any boxthorns re-introduced to

the islands by birds flying from mainland infestations

will continue to need to be controlled, although the

gradual re-establishment of native plant species in

disturbed areas may ultimately limit opportunities

for new boxthorn plants to establish. The destruction

or degeneration of habitat, or the introduction of

predators, can cause the demise of many native

species, both plants and animals. The monitoring of

introduced species, and control where necessary, will

help conserve both the native vegetation and faunas

of these islands. A systematic and sustained program

to eradicate the introduced population of house mice

from the island could be planned as part of future

conservation management.

Generally the light-station complex is in fair

condition. The light-station and other evidence of

human occupation should be conserved to protect

their cultural significance and adapted as necessary

for ongoing use. Any new development should be to

the south of the existing cottages. Regular

maintenance checks of the building should be

undertaken to mitigate the effects of weather, the

saline environment and impacts of plants and

animals. Limited access to the island by air and water

should be maintained. If possible occupancy should

be encouraged with comprehensive waste

management strategies developed.

The marine studies at Althorpe and Haystack Is

support the conclusions of earlier island studies (e.g.

the Encounter 2002 expedition; Robinson ef al.

2003) that islands support a greater diversity of

habitats than coastal locations. This is due partly to a

greater range of depths close to shore, and the higher

productivity of islands, which can capture the

production of surrounding waters and upwelling

regions (the offshore island effect). The greatest

biodiversity would be protected by maximizing the

number of habitats conserved, as shown by

Benkendorff (2005). As wave energy and light are

key forcing functions for benthic communities, a

Marine Protected Area should contain

representatives of habitats on exposed to sheltered

areas from the intertidal to deep water, embracing all

islands in the group.

Acknowledgments

In addition to the major sponsors of the expedition,

SARDI (Aquatic Sciences) and DEH, we thank Rob

Lewis and Anthony Cheshire for their strong support.

FoAICP supported the terrestrial program with a

grant of $3,520 from the Australian Government

Envirofund, sponsored by Friends of Parks Inc., and

the Department for Environment and Heritage

provided a small boat and various support staff. We

thank the Ngerin crew, and all support staff for their

contribution, and in particular Mick Clark (Dive

Supervisor), for ensuring the safe conduct of a

diverse diving program.

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symbiosis among inshore fishes at Althorpe Island,

South Australia and elsewhere. Trans. R. Soc. S. Aust.

129, 193-201.

StapLes. D. A. (2005) Pycnogonida of the Althorpe Islands,

South Australia. Trans. R. Soc. S. Aust. 129, 158-169.

WALKER-SMITH, G. K. A new species of Neopeltopsis

(Copepoda, Harpacticoida, Peltidiidae) from Althorpe

Island, South Australia. Trans. R. Soc. S. Aust. 129,

170-182.

ZANG, W-L. (2005) Geology of Althorpe Island. Trans. R.

Soc. S. Aust. 129, 90-93.

Transactions of the Royal Society of S. Aust. (2005), 129(2), 90-93.

GEOLOGY OF ALTHORPE ISLAND

by W. L. ZANG*

Summary

ZANG, W. L. (2005) Geology of Althorpe Island. Trans. R. Soc. S. Aust. 129(2), 90-93, 30 November, 2005.

Althorpe Island is located near the southern margin of the Gawler Craton and contains a Palaeoproterozoic

granite basement, which is capped by Quaternary calcarenite of the Bridgewater Formation. The basement

(Donington Suite) was multiply deformed and intruded by the Tournefort Dyke Swarm. The Bridgewater

Formation contains mainly calcarenite and palaeosol, and in the middle part, some shelly fossils occur. The

island was probably isolated from the mainland some 9-10 000 yrs ago.

Introduction

The geology of Althorpe Island is relatively well

known and the rocks on the island can be correlated

with those on Yorke Peninsula. Tectonically, the

island is located within the southern Gawler Craton,

which contains rocks ranging from ~2550 Ma

(million years before present) to ~1500 Ma. Early

geological mapping by Crawford (1965) suggested

the island contains metamorphic basement and

Quaternary sedimentary cover; the basement rocks

were tentatively assigned to the Archaean.

Subsequent surveys by Major (1973) and Rankin et

al. (1991) interpreted the basement rocks to be of

Palaeoproterozoic age. Generally the island

comprises a coastal platform of granite basement

(Donington Suite, 1850 — 1855 Ma), capped by

Quaternary Bridgewater Formation. An updated

geological map of the island and mainland has been

published recently (Zang 2003; Fig. 1).

Basement Rocks

Basement rocks comprise mainly adamellite, and

are best displayed in the southern coastal areas (Fig.

2a). The adamellite is pink to pink-brown and

contains quartz (30 — 35%), microcline (20 — 45%)

plagioclase (15 — 30%), biotite (3 — 5%), magnetite

(1 — 4%) and accessory hornblende, sphene, apatite

and zircon. An imprecise Rb-Sr date derived from

adamellite from Althorpe I. and nearby islands

suggests an age older than 1794 Ma (Webb et al.

1986). A fine-grained biotite-hornblende micro-

* Geological Survey, Department of Primary Industries

and Resources, South Australia.

Email: zang.wen-long@saugov.sa.gov.au

Purvis, A. C. (1999) Mineralogical report Nos 7770 and 7834.

Pontifex and Associates Pty Ltd. Petrographic reports

(unpublished).

Schaefer, B. F. (1998) Insights into Proterozoic tectonics from the

southern Eyre Peninsula, South Australia. Ph.D. thesis, University

of Adelaide.

adamellite also occurs on the island as isolated pods

or dykes in the host adamellite and the presence of

orthopyroxene (~2%) suggests high igneous or

metamorphic temperatures (Purvis').

The oldest rock on the island is migmatitic

paragneiss in the northern part of the island. The

paragneiss is layered with orthoclase-plagioclase-

rich bands of garnet-biotite-spinel-sillimanite-

corundum and _ plagioclase-orthopyroxene-quartz

assemblages. This was possibly derived from a

silica-deficient siltstone (Purvis'). The mineral

assemblages suggest upper amphibolite to granulite

facies metamorphism (700 — 800 °C). Similar rocks

at Corny Point, Yorke Peninsula have been dated by

U-Pb zircon geochronology: cores of zircon in

paragneiss range in age from ~2400 Ma to ~1920

Ma, suggesting a maximum age for the sediment

protoliths, and newly-grown outer zones indicate a

metamorphism age of 1845 Ma (Zang & Fanning

2001).

Dark-coloured dykes of Jussieu Metadolerite,

intruding the Donington Suite granite, occur in the

north of the island. The metadolerite is mylonitic and

contains relics of back-veining textures, in which the

mafic rock is broken into angular fragments by the

injection of felsic magma from the host rock. This

suggests that the emplacement and crystallisation of

the mafic dykes had proceeded when host rocks were

still partly molten, and therefore syntectonically

(~1850 Ma). There are also several ultra-mafic (S10,

= 39.2%, Rankin ef al. 1991) enclaves which

comprise hornblende, clinopyroxene, plagioclase and

opaques. The ultra-mafic rocks are fine to medium-

grained and granoblastic; several outcrops in the

field record the presence of an earlier fabric

(clinopyroxene — hornblende elongation).

Undeformed, younger conjugate mafic dykes

(Tournefort Dyke Swarm) trend NW-SE and NE-SW,

showing a conjugate relationship. These dykes are

generally altered and consist mainly of plagioclase,

hornblende, clinopyroxene, biotite, sphene, opaques

etc. They may be as old as ~1810 Ma (Schaefer?).

GEOLOGY OF ALTHORPE ISLAND

28,981

Ae Af

<= re

St Kilda

Formation ® Ultramafics

| Bridgewater

__| Formation 6 Microademellite

7 7) Tournefort Jussieu

7 /7\ Dyke Swarm* Metadolerite ¢

S RAS

Donington © Paragneiss 9 bi 0

Suite 3

METRES

Fig. 1. Geological map of Althorpe I.

Sedimentary Cover

Quaternary Bridgewater Formation is ~90 m thick

on Althorpe I. (Fig. 2b), and comprises a succession

of Pleistocene beach-dune calcarenite complexes

(common along coastal settings in South Australia,

Sprigg & Boutakoff 1953). Three units can be

recognised on Althorpe I. The lower unit contains

Late-Middle Pleistocene medium to coarse-grained,

large-scale bedded calcarenite, interbedded with red-

brown sandy clay (palaeosols). Wilson? suggested that

this unit could be as old as 0.7 Ma. The middle unit is

equivalent to the Glanville Formation, from which two

pulmonate gastropods were reported (Ludbrook

1973). The formation was dated at 0.1-0.13 Ma by the

Uranium, TL and Sr methods (Belperio 1995; Zang

2003). The upper unit contains fine to medium-

grained aeolianite and bioclastic calcarenite and was

dated by the '*C method at Gleesons Landing, SW

* Wilson, C. C. (1991) Geology of the Quaternary Bridgewater

Formation of southwest and central South Australia. Ph.D. thesis,

Flinders University of South Australia).

Yorke Peninsula. Charcoal from a solution hole was

dated at 22400 + 800 years and an underlying shell

horizon at >27 000 years B.P. (Wilson*). The dates

may limit the uppermost Bridgewater Formation to

~22 000 years. The Bridgewater Formation on the

island is capped by calcrete.

The calcarenites or calcareous aeolianites of the

lower Bridgewater Formation are considered to

have been deposited as a series of spatially

separated, shoreline-parallel dunes or barriers.

Morphologically, individual dunes display weakly

lenticular to tabular geometries and are capped by a

red-brown palaeosol layer. Generally, the

calcarenites consist of homogeneous, fine to

medium-grained, well-sorted bioclastic grainstone

with tiny fragments of molluscan, bryozoan,

foraminiferal, echinoid, algal, peloidal and lithoclast

allochems (Wilson’).

Near the jetty, modern beach sand (St Kilda

Formation), mostly silica sand with some carbonate

shells, has been deposited. This beach provides the

only safe boat-landing spot on the island.

The top of the island is capped by a thin layer of

W. L. ZANG

Fig. 2. A) basement granite of the Donington Suite, Mooring Bay, Althorpe 1. (1950-1855 Ma); B) Quaternary sediments

of the Bridgewater Formation, Mooring Bay, Althorpe I. (~0.5 Ma).

GEOLOGY OF ALTHORPE ISLAND 93

calcrete. Calcrete and soil weathering profiles are

extensively developed within the calcareous

aeolianites and play an important role in the

stabilisation and preservation of these sediments.

Calcretisation was obviously developed during

several pedogenic stages of biochemical and

physiochemical processes to a completely indurated

or fossilised profile. Calcrete was used for building

material in the early years and is still mined for road

building material on Yorke Peninsula.

Palaeogeography

Althorpe I. and the surrounding isles are

considered to have been linked with Yorke Peninsula

during deposition of the Bridgewater Formation ca.

half a million years ago. Repetitive transgressions of

the sea across the continental shelf caused successive

calcarenite barriers to form (Belperio 1995). Climate

and sea level change were driven by repetitive global

Quaternary glaciations and deglaciations. Complete

separation of the islands from the mainland by gulf

waters may have occurred as recently as 9-10 000 yrs

ago when the land bridge was eroded and submerged

during the latest sea level rise. Some possible

marsupial bones in shallow caves on the island may

support this suggestion.

Discussion

The geological history of the Althorpe Is can be

traced back to ~2000 Ma years ago when the region

of Yorke and Eyre Peninsulas was a palaeo-ocean

bounded by the Archaean terrains of the northern

Gawler Craton and southern Antarctic continent.

Remnants of metamorphosed siliciclastic and

carbonate deposits on the island and Yorke Peninsula

may suggest a shallow marine environment in the

areas (Zang & Fanning 2001). Those shallow marine

deposits were intruded by granites of the Donington

Suite during ~1850 - 1855 Ma and metamorphosed

by deep-crust sourced heat, up to ~800 °C. The

granites, in return, were intruded by mafic

Tournefort Dyke Swarm during ~1800 Ma. No

geological history has been recorded on the island

until the middle Quaternary.

Middle Pleistocene (~0.5 Ma) deposits of the lower

Bridgewater Formation in the region might be related

to a short glacier meltdown and deposition of

regressive calcarenites, of which the uppermost is

always weathered or veneered by palaeosol. The

glacial-transgression deposits of the Upper

Pleistocene Glanville Formation reached the

Althorpe Is, and as far as the southern margin of

Yorke Peninsula. A Late Pleistocene uplift event is

recognised by the formation of Peesey Swamp to the

east of Warooka township, Yorke Peninsula, where

the Middle Pleistocene Bridgewater Formation

forms the ridge to the west of the Warooka Fault

zone, The Glanville Formation and overlying

aeolianites were deposited in the Peesey Swamp

depression. Related to this uplift, the uppermost

Bridgewater Formation in the areas contains mainly

palaeosol and weathered calcareous sand.

The Holocene St Kilda Formation is probably

deposited following more recent — glacial

transgression, starting some 6000-7000 years ago in

South Australia (Belperio 1995). This transgression

reached southern Yorke Peninsula, since Posidonia-

rich deposits occur in the Marion Lake area (von der

Borch et al. 1977). The relative sea level in the

Althorpe Is and southern Yorke Peninsula areas

seems to have risen slightly during the last 6000

years.

References

Bevcrerio, A. P. (compiler) (1995) Quaternary. /n: Drexel,

J. F & Preiss, W. V. (Eds), The geology of South

Australia, Vol. 2: The Phanerozoic. S. A. Geological

Survey Bulletin 54, 219-281.

Crawrorb, A. R. (1965) The geology of Yorke Peninsula.

Geological Survey. S. A. Department of Primary

Industries and Resources. Bulletin 39, 1-96.

Lupsrook, N. H. (1973) Two pulmonate gastropods from

Bridgewater Formation, Althorpe Island. S.A.

Department of Mines. Report Book 73/00206.

Major, R. B. (1973) Preliminary report — geology of

islands of the western continental shelf of South

Australia. S. 4. Department of Mines. Report Book

73/226.

RANKIN, L. R., FLINT, R. B. & BELPERIO, A. P. (1991)

Precambrian geology of islands in the Investigator Strait

area, South Australia. S. A. Department of Mines and

Energy. Report Book 91/57,

Sprica, R. C. & BouTakorr, N, (1953) Summary report on

the petroleum possibilities of the Gambier Sunklands.

Mining Review, Adelaide 95, 41-62.

VON DER Borcu, C. C., BOLTON, B. & WARREN, J. K. (1977)

Environmental setting and microstructure of subfossil

lithified stromatolites associated with evaporites. Marion

Lake, South Australia. Sedimentology 24, 693-708.

Wess, A. W., THOMSON, B. P., BLissetTT, A. H., DALY, S. J.,

FLINT, R. B. & Parker, A. J. (1986) Geochronology of

the Gawler Craton, South Australia. Aust. J Earth

Sciences 33,119-143.

ZANG, W. L. (2003) Maitland Special Map Sheet. S. A.

Geological Survey. Geological Atlas 1:250 000 Series,

sheet S153-12 and portion SI53-16.

& FANNING. C. M. (2001) Age of the Kimban

Orogeny revealed — U-Pb dates on the Corny Point

Paragneiss, Yorke Peninsula, South Australia. MESA

Journal 23, 28-33.

Transactions of the Royal Society ofS. Aust. (2005), 129(2), 94-99.

HUMAN SETTLEMENT ON ALTHORPE ISLAND AND

CONDITION OF THE LIGHTHOUSE COMPLEX

by A. RADFoRD!

Summary

Raprorp, A. (2005). Human settlement on Althorpe Island and condition of the lighthouse complex. Trans. R.

Soc. S. Aust, 129(2), 94-99, 30 November, 2005.

Althorpe I. became important for coastal navigation with construction of the lighthouse in 1879. The island

was occupied from then until 1991 when the light was automated. Evidence of human occupation during that

period included: various buildings associated with the lighthouse, runway, jetty and flying fox, grave sites and

guano mining. The present significance of the island in terms of its cultural heritage, and other values is

discussed.

Key Woros: Althorpe I., history, lighthouse, jetty, flying fox, exploration, shipwrecks, South Australia.

Introduction

The Althorpe Is are located 8 km from Cape

Spencer at the southern tip of Yorke Peninsula.

During the period of European exploration the island

was first visited by Matthew Flinders (Cooper 1953).

Louis de Freycinet, cartographer to Nicholas

Baudin’s French expedition, named the island group

“Iles Vauban” (Brown 2004) and the main island Ie

Laubadére (Péron 1824). About 10 years later, when

Flinders drew up his maps, the island and the nearby

Western Isles were named Althorpe? Islands. They

were named after Viscount of Althorp, son of Earl

George John Spencer (Farlex Inc. 2005)>. Earl

George Spencer, 2nd Earl Spencer, was then First

Lord of the Admiralty and Spencer Gulf and Cape

Spencer were named after him.

The main island is a calcarenite plateau with cliffs

~90 m high over a granitic base (Zang 2005), and

provides limited opportunities for access. The beach

at Mooring Bay on the northern side was used to

construct a jetty and obtain access. Althorpe I.

became an important part of South Australia’s coastal

navigation with the development of a lighthouse and

associated structures in 1879.

This paper provides a brief historical review of the

European occupation of Althorpe I., lists the

shipwrecks that have occurred, and describes the

present state of the island’s various structures, which

comprise its heritage values.

Heritage Branch, Department for Environment and Heritage,

GPO Box 1047, Adelaide, SA 5001.

Email: Radford.Alison@saugov.sa.gov.au

The ’e was probably a spelling mistake.

The courtesy title of the eldest son of the Earl Spencer is

Viscount Althorp.

Conservation Plan for Althorpe Island Lightstation (1991)

Australian Construction Services.

Methods

Historical sources were examined and combined

with a site visit on 2 — 5 Feb. 2004, as part of the

2004 expedition (Murray-Jones & Shepherd 2005),

in order to: (1) update the 1991 Conservation Plan‘;

(2) provide a dilapidation report on the state of the

lighthouse and its precincts, keepers’ cottages,

associated sheds, and the jetty precinct; and (3)

prepare policies for the future of the built form and

shipwrecks in the area. Other evidence of human

occupation including guano mining, grave sites, and

associated lighthouse keeper activities were also

noted. Sites were visited, GPS coordinates taken

(AMG Coordinates using Datum WGS84), measured

drawings undertaken of the lighthouse precinct, and

a condition audit conducted. The location of places

or sites referred to are shown in Murray-Jones &

Shepherd (2005).

Results

History

Early settlement to South Australia was by sea to

Port Adelaide with an uncertain route through

islands, reefs and strong tidal currents. Light towers,

such as pharos, were recognised as valuable aids to

help ships navigate through these dangerous waters.

The British Trinity House was set up Royal Charter

in 1514 to serve as a charitable organisation for

mariners, provide aids to navigation and to care for

lighthouses (Trinity House 2005). Trinity House in

Port Adelaide was founded for the same purposes on

the British model.

The first lighthouse in South Australia was built at

Cape Willoughby, on Kangaroo I., and comm-

issioned on 10 Jan. 1852. Others followed in the

vicinity. Following an Intercolonial Conference in

HUMAN SETTLEMENT 95

1873, a further lighthouse construction program

began. One such proposed lighthouse location was

Cape Spencer, but after Commander Goodbrough

visited Spencer Gulf in 1874 he recommended

establishing a light on Althorpe I. instead.

In 1876 estimates in the Public Services Loan Bill

provided £6,480 to construct a lighthouse and

lightkeepers’ cottages on the island. After some

delays, drawings were finally prepared by Robert

Hickson, Engineer of Harbors and Jetties, South

Australia in 1877, but there were further delays

through lack of tenders. As a result, construction was

finally undertaken by Harbors and Jetties staff in

1878, but not without accident®.

Due to the steep cliffs, a jetty and an inclined

ladder were constructed first. The attending cutter,

Young St George, was wrecked and later, the

foreman, John Anley, was killed by a falling rock.

Costs escalated, due inter alia to the need to

transport all the drinking water from Port Adelaide,

and from delays in obtaining the Chance Brothers

revolving light. The light was finally exhibited on

14 Feb. 1879 at a total cost of £11,700.

Over the next century various repairs and changes

were carried out including additional water storage

(an ongoing problem for lightstation staff). A

telephone line between the Island and Cape Spencer

was laid in March 1886, following fears of a Russian

invasion. The inclined ladder and tramway was

replaced with a flying fox in 1904.

The Commonwealth took over management of

Australian lighthouses from Trinity House in 1912,

and the late 1930s to early 1940s saw a major

upgrading program for the site. This included new

walls, a new engine house and signal box, upgrading

of the cottages to provide new bathrooms, rainwater

> Ibid,

1 Thid.

* Heritage Branch files — Statements of Heritage value:

Lighthouse (10312): Completed in 1879, the Althorpe Island

Lighthouse is significant for its association with the establishment

of the network of lighthouses around South Australia’s shores to

protect the shipping routes, which were crucial to the State’s

economic development at the time. The lighthouse is important as

an example of a well-executed stone lighthouse tower, in

particular the detailing of the spiral staircase (HAS 2/2000).

Keepers’ cottages (10314); The three lighthouse cottages were

constructed to service the Althorpe Island Lighthouse and remain

as an intact example of lighthouse keepers’ cottages of the late

1870s. They are an integral part of this remote lightstation, which

was established to guide both local and interstate shipping through

Investigator Strait during the period when sea transport was vital

to the State’s economy (HAS 2/2000).

Jetty (10318): This jetty and associated structures were built to

service the Althorpe Island lightstation, established in 1879 to

protect the shipping routes which were crucial to the State’s

economic development at the time. The jetty represents the

dependence on maritime transport as the only means of servicing

the State’s island-based lightstations and reinforces the difficulties

posed by the remoteness of these sites (HAS 2/2000).

” Register of National Estate Identified No. 6887.

SA Government Gazette 16 March 1967.

'! SA Government Gazette 14 August 1997,

storage, and replacement of the rear pitch of the main

roof and lean-to with a single pitch. The main light

was converted to electricity in 1963 and a 3 m

leading light erected in 1963.

During the programme of automation of Australian

lighthouses from 1975, the Althorpe I. lighthouse

was finally automated in 1991, and the island

vacated by the Australian Maritime Safety Authority

(AMSA)’. The lighthouse and other built structures

became part of the Althorpe Islands Conservation

Park and were occupied occasionally.

The lighthouse (10312, Fig. 1), keepers’ cottages

(10314, Figs. 2, 3) and jetty (10318, Figs. 4, 5) were

listed in the State Heritage Register in 1980, and

Statements of Heritage prepared’.

The lighthouse and cottages were also listed on the

complex is located within the Unincorporated area of

the State.

The Western Isles above high water were gazetted

as a Fauna Reserve in 1967!° and became the

Althorpe Islands Conservation Park in 1972.

Haystack and Seal Islands were added in 1977 with

the area below high water added in 1991. Althorpe I.

(Sections 13 and 61) was added to the Conservation

Park in 1997 (Robinson et al. 1996)!'. The

Conservation Park is important for its cultural and

natural features, and as a habitat for sea birds.

The Buildings

The lighthouse precinct (Fig. 1) consists of a

lighthouse, three keepers’ cottages, various sheds,

tanks, fences and paths and the remnants of a

runway.

The lighthouse (Fig. 3) (Lat. 35° 22.2’ S; Long.

136° 51.7’ E) is managed by AMSA. It is a circular

13 m tall structure constructed of random rubble

limestone and painted white. The exterior was altered

in 1973 with the sandstone quoins being replaced

with concrete. The light, exhibited at 107 m above

sea level, is a 120 volt 1000 W tungsten halogen

lamp with a range of ~45 km.

The interior consists of a spiral stair of cut

cantilevered sandstone risers with treads finished in

Mintaro slate (now painted alternately red and

green). The balusters are steel with an unpainted

handrail. The floors are Mintaro slate supported by

cast iron joists.

The cottages all show a typically South Australian

design unlike many lighthouse complexes elsewhere

where there is a hierarchy of living conditions based

on rank, Cottage | has a window from the master

bedroom towards the lighthouse (absent in Cottages

2 and 3). The three cottages have received basic

maintenance only, since the lighthouse was

automated, and all show evidence of the harsh marine

climate.

96 A. RADFORD

Legend

. Cottage (Principal keeper’s residence)

. Cottage (2nd keeper’s residence)

. Lighthouse

. Generator Shed

. Workshop

Office

Rope Locker

CIDARWNE

. Cottage (Ist assistant keeper’s residence)

Fig. 1. Lighthouse precinct plan (adapted from Conservation Plan).

Jetty precincts

The jetty area consists of the flying fox (Fig. 4),

jetty (Fig. 5), shed and nearby grave-site for the

bodies recovered from the SS Pareora (Fig. 6). The

jetty is a wooden structure of varying age, 72 m long

and 2.7 m wide, with a hand crane at the seaward end

standing on a wider 3.6 m platform. The jetty is now

in poor condition with many structural members

corroded, eaten away, missing, or, in the case of piles,

replaced with newer, round, sister piles bolted beside

them". A flying fox attached to the jetty terminates

'2 National Archives have reference to repairs at various times but

they are closed files.

'3 Lighthouse log 2 May 1892 notes “employed putting up a fence

around the grave”. National Archives, Adelaide Series D19

barcode 10684806. Unfortunately the logs for the time of death

are missing.

at the top of the cliff. While the top anchorage is in

good condition, the flying fox is deemed unsafe due

to the poor condition of the lower anchorage to the

jetty.

Graves

There is much evidence of the 127-year human

occupation all over the island. This includes graffiti

in Cathedral Cave on The Hump, and the remains of

a stone wall, which may have been a barrier to

prevent animals crossing from the main island on to

the Hump.

Four graves can be seen above Mooring Bay. They

may be memorials or may have been modified and

updated by lighthouse staff and other island

occupants over time. These are:

(a) Julies Garbis, located near top of slope to E of

HUMAN SETTLEMENT 97

Bedroom

Enclosed veranda |

es =

0 3000mm

eel

Bedroom

iain |

Fig. 2. Typical cottage plan (from Conservation Plan),

Fig. 3. Cottage with lighthouse at rear.

jetty. The grave is bounded with old bed posts'3,

with wording on the grave ‘Julies Garbis died May

8th 1890 aged 42 yrs’.

(b) Near sea level, to the east of the jetty, is located

a white timber cross with a lighthouse carved on the

top in a pile of rocks. It has no inscription but may

be in memory of Arthur (‘Dick’) Johnson, a crew

‘4 Memorandum from Head Keeper to District Office Port Adelaide

11 November 1919. National Archives, Adelaide Series D14

barcode 430591.

Fig. 4. Flying fox from jetty.

Fig. 5. Jetty from above.

member who lost his life on the wrecked cutter

Rapid. His body was never found (Coroneos &

McKinnon 1997),

fenced area'* with a cross (Fig. 6) inscribed: ‘SS

Pereora (sic) Wrecked 18.9.1919 JF Booth JC

Braithwaite R Deebly RIP’.

(d) Further west at the base of a cliff is a replica

marker — an old door inscribed: ‘To the memory of

G Peterson aged 48 yr died October 8th 1838’.

Guano mining and sundry remains

Penguins inhabit many limestone caves around the

island, including some close to the cliff top. These

98 A. RADFORD

Fig. 6. Pareora grave.

caves often have low head room (<I m) and

penetrate some metres into the cliffs. The lighthouse

logs mention boats arriving and collecting guano'’,

and there is evidence in some of the caves of

abandoned clothing and equipment suggesting

guano mining activities.

On the western side of the island, at sea level, are

some rusty metallic remains of old island machinery

and possible relics from the Altair wreck (July

1971).

Sealing

It is known from historical records that seals were

hunted for their skins on many of South Australia’s

offshore islands, including the Althorpe Is. The

earliest reference is in 1815 when crew of the brig,

Spring, reported sealing around Kangaroo I. and the

Althorpes (Sexton 1990). As late as 1892 the crew

of the wrecked cutter Welling were assisted by a

party of sealers operating at the island (Coroneos &

McKinnon 1997). Thus far, no physical evidence of

this activity has been located on Althorpe I.

Various references in lighthouse logs 1882-1892. National

Archives, Adelaide Series D19.

’ Various telegrams from Head Keeper to District Office, Port

Adelaide 18 September 1919 National Archives Series D14

barcode 430591,

Shipwrecks

The six recorded shipwrecks adjacent to Althorpe I.

all happened during or after construction of the

lightstation (Arnott 1996). Three have significance

in being wrecked while the lightstation construction

was under way or because of the loss of life. The

details given below are extracted from Coroneos &

McKinnon (1997).

(a) Young St George was wrecked on 3 Jan. 1878

during construction of the lightstation. The yacht was

built in Melbourne in 1863, and later sold to the SA

Government, sometime after 1873, and used as a work

vessel. The vessel parted from its moorings during the

night and drifted on to rocks in Mooring Bay. The

crew managed to reach safety and next morning took

everything moveable from the wreck. The hull was a

complete loss, and has not been located.

(b) SS Pareora was wrecked on 18 Sept. 1919 due

to poor navigation, and ranks as one of the worst

disasters in Investigator Strait. The screw steamer,

first named Breeze, was built in Newcastle-upon-

Tyne, UK in 1896, and renamed Pareora in 1900

after purchase by the New Zealand Shipping

Company. A few months before its loss, the Pareora

underwent an overall refit for its new owner the

Electrolytic Zinc Company of Australia for

transporting zinc concentrates to Hobart and

returning with timber. The vessel grounded at

3.45 am and broke up during heavy seas over the next

three hours. Only seven of the 18 crew were rescued

and three bodies recovered'®. The wreck site in NW

Bay has been surveyed by the Department for

Environment and Heritage (DEH).

While sheltering from a storm at the main island, the

fishing cutter broke from its anchors and struck

rocks. After taking water it was abandoned by its

crew and subsequently lost near the Island. The

wreck has not been located.

(d) The fishing cutter Rapid was wrecked on 18

Sept. 1937, while sheltering with other vessels in

Mooring Bay during a storm. She lost her anchor at

3 am, drifted ashore to the east of the jetty and broke

up on the rocks. The owner, his wife and one of the

crew managed to reach shore, but crewman, Arthur

(‘Dick’) Johnson disappeared. The wreck was

washed up on shore the next morning. The wreck has

not been found.

(e) The fishing cutter A/tair was wrecked in July

1971. Little is known about the loss of the cutter

other than it ran aground near The Monuments at the

NW end of Mooring Bay. Wreckage, believed to be

remains of the vessel, has been located.

(f) The shark fishing vessel Mylor Star was

wrecked during a storm on 9 June 1982, after being

driven ashore on the south side of Althorpe I. The

location of the wreck is known.

HUMAN SETTLEMENT 99

Discussion

The built structures on the island are currently in

good to fair condition, but lack of use and regular

maintenance have allowed vegetation and shearwater

nests to encroach, and the extreme weather

conditions are taking their toll. The runway is

undermined by shearwater nests and is no longer

usable. Some outhouses and fences are in a state of

disrepair.

The lighthouse infrastructure is an important

record of a past era of South Australia’s history. The

lighthouse, maintained by AMSA, is now automatic

and the main interest in the islands is now focused on

their terrestrial and marine natural history. The island

infrastructure of cottages, jetty and flying fox is

maintained for that purpose, largely by a dedicated

group (Friends of the Althorpes Is. Conservation

Park), which includes previous residents.

General future policies from the Conservation Plan

relate to the Burra Charter principles of adaptation,

preservation, conservation and restoration (Walker &

Marquis-Kyle 2004). The Conservation Plan

recommends retention of the three cottages,

lighthouse and leading light with adaptation for

ongoing use. Other buildings on the island do not

require conservation, but their removal should

minimise impacts on future use of the site. Any built

form to be removed should be recorded (graphically

and photographically) and carefully dismantled with

removal of any material that may blow around or

cause future pollution. The Conservation Plan also

recommends that retention of the flying fox should

be encouraged and the jetty should be preserved,

though it does recognise this may only be for the

short term!7,

New development and adaptation is covered in the

Conservation Plan. The lighthouse should remain the

dominant element and the cottages providing a lineal

alignment with no development to the north'®.

Acknowledgments

I thank RV Ngerin and crew for transportation to

Althorpe I. and the Department for Environment and

Heritage for its support of the terrestrial program.

Particular thanks to Vicki Cheshire for her assistance

in surveying the lighthouse precincts, to Erika

Lawley for her historical research and for pointing

out important features on the island, and to

anonymous referees and editors for improvements to

the manuscript.

References

Arnott, T. (Ed.) (1996) “Investigator Strait Maritime

Heritage Trail”. (DENR, Adelaide).

Brown, A. J. (2004) “Ill Starred Captains: Flinders and

Baudin”. Revised ed. (Fremantle Arts Centre Press,

Perth).

Cooper, H. (1953) “The Unknown Coast; being the

exploration of Captain Flinders RN along the shores of

South Australia 1802”. (Advertiser Press, Adelaide) cited

in ACS (1991) Conservation Plan for Althorpe Island

LightStation.

Coroneos, C. & McKINNon, R. (1997) “Shipwrecks of

Investigator Strait and the Lower Yorke Peninsula”.

South Australian Maritime Heritage Series No.4 (DENR,

Adelaide).

FARLEX INC. (2005) The free dictionary. Accessed 26 Aug.

2005. URL http://encyclopedia.thefreedictionary.com

Murray-Jones, S. & SHEPHERD, S. A. (2005) Expedition to

the Althorpe Islands, South Australia: introductory

narrative, and marine conservation recommendations.

Trans. R. Soc. S. Aust. 129, 85-89.

'7 ACS (1991) Conservation Plan for Althorpe Island Light Station

— Section 5

'S Ibid. — Section 6.3

Peron, F. (1824) “Voyage of Discovery to the Southern

Lands”. 2nd Ed. 1824 translated by Christine Cornell

(2003), (The Friends of the State Library of South

Australia, Adelaide).

Rosinson, A. C., CANTy, P. D., Mooney, P. & Rupbuck, P.

(1996) “South Australia’s Offshore Islands” (Australian

Government Publishing Service, Canberra).

SexTon, R. T. (1990) “Shipping Arrivals and Departures

South Australia 1627-1850”. (Gould Books — Roebuck

Society, Adelaide).

Trinity House (2005) Trinity House Lighthouse Service.

Accessed 26 Aug. 2005. URL http://www.trinityhouse.

co.uk/

WaLKer, M. & MARQUIS-KYLE, P. (2004) “The Illustrated

Burra Charter: Good Practice for Heritage Places,

Australia.” (ICOMOS Inc, Burwood).

ZANG, W-L. (2005) Geology of Althorpe Island. Trans. R.

Soc. S. Aust. 129, 90-93.

Transactions of the Royal Society of S. Aust. (2005), 129(2), 100-110.

LAND USE AND VEGETATION OF ALTHORPE ISLAND, SOUTH AUSTRALIA,

AND A FLORISTIC COMPARISON WITH SOUTH NEPTUNE ISLANDS

by E. F. LAWLEY! & S. A. SHEPHERD?

Summary

LawLey, E. E. & SHEPHERD, S. A. (2005). Land use and vegetation of Althorpe Island, South Australia, and a

floristic comparison with South Neptune Islands. Trans. R. Soc. S. Aust. 129(2), 100-1 10, 30 November, 2005.

A brief history of the land use of Althorpe Island, during its occupation from 1879 to 1991, is presented and

the effects of occupation on the vegetation are described. In all, 46 native and 40 introduced plant species are

listed, together with a species list from the South Neptune Islands for comparison. Six vegetation communities

are described and a revised vegetation map of the island is provided, based on a floristic analysis of 10 vegetation

quadrats, and interpretation of aerial photography. The likely extinction of some species and vegetation changes

following extermination of goats and sheep from the island are discussed.

Key Worpbs: Island flora; vegetation; plant species; photo points; lighthouse station; Althorpe Island; South

Neptune Islands; Puffinus tenuirostris; Short-tailed shearwater; goats; exotic plants; introduced species.

Introduction

The flora of offshore islands often has unique

characteristics because islands are exposed to

patterns of colonization and extinction that depend

on distance from sources of colonists, size of the

islands, and time of isolation from the mainland

(MacArthur & Wilson 1967). Further, historical

events, such as disturbances, exotic species and

grazing, and local climate may affect the vegetation

of islands in different ways than on the mainland

(reviewed by Brown & Lomolino 2000). The

Althorpe Islands (Latitude 35° 23'; Longitude 136°

51’), taken here to refer to Althorpe I. and the nearby

Western Isles, are 8.5 km from the southern tip of

Yorke Peninsula. The main island, Althorpe I. is an

elevated plateau covering 92 ha and is 93 m above

sea level, with a smaller plateau, the Hump, attached

by a depressed nexus called the Saddle (Murray-

Jones & Shepherd 2005). The average annual rainfall

is 512 mm. The maximum depth separating the

Althorpe Is from the mainland is about 25 m, so they

would have become isolated when sea-level rose ~9-

10 000 years ago (Belperio et al. 1983). Althorpe I.

is unusual because it has been grazed, and the topsoil

extensively burrowed by a large population of

nesting Short-tailed shearwaters (Puffinus

tenuirostris) (Robinson et al. 1996), and to a small

extent by the Little penguin (Eudyptula minor).

' PO Box 89, Tarlee, South Australia 5411.

Email: lawley@chariot.net.au

> SARDI Aquatic Sciences, PO Box 120, Henley Beach, South

Australia 5022.

‘Unpublished report by Graham Medlin for Friends of Althorpe

Islands Conservation Park. 2001.

4SA Lighthouse logs. NAA: D19,1908-1910, in National Archives

of Australia.

° SA Lighthouse logs. NAA: D19, 1879-1880, 1883-1888.

A description of the vegetation and an

accompanying floristic map were first published by

Robinson er a/. (1996), following a visit in 1982.

This paper summarises the history of land use of

Althorpe I., describes its vegetation and the major

biotic influences on the vegetation, and gives a plant

species list (Table 1). For comparison, we also

provide a plant species list from two neighbouring

islands of similar size to Althorpe I., the South

Neptune Islands, here termed the SN Lighthouse I.,

and SN North I. The former of these has been

similarly grazed, but the latter has had no significant

history of any human impact.

Human impact after 1879

Nothing is known of the native flora or fauna

present on Althorpe I. in 1879, when the lighthouse

station was built and thereafter occupied by three

families. Sub-fossil remains found in 2001 on, or just

under, the surface in caves on the island indicate the

relatively recent extinction of the Greater sticknest

rat (Leporillus conditor), the Bush rat (Rattus

fuscipes), and the Southern brown bandicoot

(Isoodon obesulus?).

Goats were probably introduced soon after human

occupation, and by 1908 they were being hunted. By

1962 there were 120 goats, but a major cull in 1968

left only six, which bred up to 25 by 1981. By about

1990, however, all remaining goats had been

exterminated.

Other domestic grazers, such as a horse and groups

of 30-40 sheep at a time, were introduced at various

times‘, and may have affected the vegetation directly

by grazing, and indirectly by introducing seed

variously present in their gut or wool on arrival on the

island. Firewood, which was regularly brought to the

island, is another likely source of introduced seed.

VEGETATION OF THE ALTHORPE ISLANDS 101

Cleared or Non-Native Grasslands

Atriplex paludosa ssp. cordata

Low Open Shrubland

Myoporum insulare

Low Open Shrubland

Maireana oppositifolia | Atriplex

paludosa ssp. cordata

Low Shrubland

Nitraria billardierei

Low Shrubland

Halosarcia spp.

Low Closed Shrubland

Bare Rock or Sand

St Survey Sites Ca,

668500

6085000

Cee

6084500

6084000

6083500

Geocentric Datum of Australia, 1994

@ |e

and Hetilage

669000

Fig. 1. Floristic map of Althorpe Island showing all vegetation communities and the location of all survey quadrats, ALT

00101-ALT 010001.

The vegetation was also affected through

collection of building and paving materials®. Sand

was removed from an area of 600 m2 to the west of

the lighthouse, called the Quarry, which in 1980 was

still almost without vegetation. “Seed grass”, couch

grass (Cynodon dactylon), and marram grass

® SA Lighthouse logs. NAA: D19, 1881-1883.

7 SA Lighthouse logs. NAA: D19, 1908-1910; 1910-1912.

* The specimens collected during 2000 were duplicated during the

2001 survey.

(Ammophila arenaria), were introduced in 1910 and

1911’, but the marram grass did not become

established (Table 1). In 1961, an airstrip, 450 m

long, was cleared on the plateau, and sown with

mixed grass seed for surface stabilization.

Prior to this study, plant collections on Althorpe I.

were made in 1907 by Dr R. S. Rogers (Maiden

1908), in 1910 and 1916 by Captain S. A. White

(White 1916), in 1982 by Robinson ef a/. (1996), and

in 2000 by S. Hawkins*.

102 E. F LAWLEY & S. A. SHEPHERD

TABLE |. Plant species recorded from Althorpe Island and the two South Neptune Islands. Taxonomy follows the Census of

South Australian Vascular Plants. Ed 5.1 (May 2004). Introduced species are marked with an asterisk. Species lists

presented are the result of historical records and terrestrial surveys in 1982 and 2001.

New records for Althorpe I. collected from 2001-2004 are marked as +.

The 1982 survey of South Neptune Islands by Robinson et al. (1996) is omitted as it duplicated the list of plant species

occurring on SN North I. found by Stirling et al. (1970).

The Stirling 1970 list is included because some species not found in 2001, were likely present, but not all collection areas

on SN North I. were revisited, due to time constraints. SNI Lighthouse I. was thoroughly searched over 10 days resulting

in a comprehensive species list.

2 s ¢

2 =e =e & @

ss ss s & a 4 va

Scientific Name Common Name mn & - a & 8 &

Ba B@ B &B B = 3

Pp ege oF? & @

an ia.” aa

3 3 I

A A A

year of collection 1907 1916 1982 2001 2001 1970 2001

CASUARINACEAE

Allocasuarina verticillata Drooping sheoak *

(.keeper intr c.1982)

URTICACEAE

Parietaria debilis Smooth nettle * *

Urtica urens* Stinging nettle *

POLYGONACEAE

Muehlenbeckia gunnii Coastal climbing lignum * * *

AIZOACEAE

Carpobrotus edulis* Hottentot fig * *

Carpobrotus rossii Karkalla % * * * *

Disphyma crassifolium Round-leaf pigface * * *

ssp. clavellatum

Mesembryanthemum Common iceplant * * *

crystallinum*

Tetragonia implexicoma Bower spinach * * * *

PORTULACACEAE

Calandrinia calyptrata Small-leaved parakeelya *

Calandrinia granulifera Pigmy purslane * *

CARYOPHYLLACEAE

Polycarpon tetraphyllum* Four-leaf allseed *

Sagina apetala* Common pearlwort * * *

Scleranthus pungens Prickly knawel * *

Silene nocturna* Mediterranean catchfly *

Spergula arvensis* Corn spurrey *

Spergularia marina Salt sand-spurrey * *

Spergularia media* Coast sand-spurrey * * * *

Spergularia rubra* Red sand-spurrey *

Stellaria media* Chickweed * *

CHENOPODIACEAE

Atriplex cinerea Grey saltbush * * * * *

Atriplex muelleri(?) Mueller's saltbush *

Atriplex paludosa ssp. cordata Marsh saltbush * x *

Atriplex semibaccata Berry saltbush *

Chenopodium murale* Nettle-leaved goosefoot * *

* * od * *

Enchylaena tomentosa

var. tomentosa

Ruby saltbush

TABLE 1. Cont.

VEGETATION OF THE ALTHORPE ISLANDS

s Ss a

ih Ee Ea

& az} a aa a Zz z

Scientific Name Common Name 8 4 4 4 FA FA g

sD) i) Oo vo i= s g

& & B 6: @ 2 @

| £2 2h $& G

= =< @ 2 — 2 &

| S 3

A A B

year of collection 1907 1916 1970 2001

Halosarcia Grey samphire *

halocnemoides ssp.

halocnemoides

Halosarcia pergranulata Samphire

Halosarcia sp. Samphire

Maireana oppositifolia Salt bluebush

Rhagodia candolleana Seaberry saltbush *

ssp. candolleana

Sarcocornia blackiana Samphire *

Sarcocornia quinqueflora Beaded glasswort

Sclerolaena uniflora Bassia

Suaeda australis Austral seablite *

Threlkeldia diffusa Coast bone fruit *

CRUCIFERAE

Brassica tournefortii* Wild turnip

Cakile maritima ssp. Two-horned sea rocket

maritima*

Hymenolobus Oval purse

procumbens*

Lepidium foliosum Peppercress * = *

Sisymbrium orientale* Wild mustard

CRASSULACEAE

Crassula closiana Stalked crassula

Crassula decumbens Spreading crassula

var. decumbens

Crassula sieberana ssp. Australian stonecrop

tetramera

LEGUMINOSAE

Medicago polymorpha var. Burr-medic

polymorpha*

Melilotus indica* King Island melilot

Swainsona lessertiifolia Coast swainson-pea *

Trifolium repens* White clover

OXALIDACEAE

Oxalis pes-caprae* Soursob

GERANIACEAE

Geranium solanderi Native geranium

Pelargonium australe Austral Stork's bill *

ZYGOPHYLLACEAE

Nitraria billardierei Nitre bush

Zygophyllum billardierei Coast twin leaf

EUPHORBIACEAE

Euphorbia paralias* Sea spurge

RUTACEAE

Common correa “

Correa reflexa var. coriacea

104

TABLE 1. Cont.

E. F LAWLEY & S. A. SHEPHERD

Scientific Name

Common Name

Althorpe Island

South Neptunes North

South Neptunes North |

year of collection 1970 2001

MALVACEAE i

Malva behriana Australian hollyhock * *

Malva dendromorpha* Tree mallow %

Malva parviflora* Small-flower marshmallow *

FRANKENIACEAE

Frankenia pauciflora Southern sea heath * *; ,

var. fruticulosa

UMBELLIFERAE

Apium prostratum Sea celery +7 * *

var. filiforme

Petroselinum crispum* Parsley *

PRIMULACEAE

Anagallis arvensis* Blue pimpernel *

Samolus repens Creeping brookweed "

LABIATAE

Prostanthera serpyllifolia Small-leaved mintbush

ssp. serpyllifolia

SOLANACEAE

Lycium ferocissimum* African boxthorn *

MYOPORACEAE

Myoporum insulare Common boobialla a

PLANTAGINACEAE

Plantago hispida Hairy plantain *

GOODENIACEAE

Goodenia varia Sticky goodenia *

COMPOSITAE

Angianthus preissianus Common cup-flower

Arctotheca calendula* Cape weed i

Brachyscome exilis Slender daisy - *

Conyza bonariensis* Flaxleaf fleabane +

Cotula coronopifolia* Waterbuttons *

Cotula vulgaris Slender cotula *

Craspedia variabilis Billy-buttons +

Euchiton sphaericus Cudweed y

Ixodia achillaeoides ssp. Coast ixodia *

achillaeoides

Lactuca_ serriola* Prickly lettuce *

Leiocarpa supina Coast plover-daisy *

Leucophyta brownii Coast cushion bush * *

Microseris lanceolata Yam daisy *

Olearia axillaris Coast daisy-bush

Olearia ramulosa * *

Podolepis rugata

var. littoralis

Pseudognaphalium luteoalbum

Twiggy-daisy bush

Pleated podolepis

Cudweed

VEGETATION OF THE ALTHORPE ISLANDS 105

TABLE |. Cont.

3 ‘

3 sS ss

g 2 @ 2 @ 5 5

Bas) & i & 4 Zz Zz

Scientific Name Common Name ne a 2 a 2 2 A

i) cP) oO o fc < =I

e a & & 2 r= B

|: 2@ BS &

= <= @ = @ 4 G&G

3 3 s

A A BAB

year of collection 1907 1916 1982 2001 2001 1970 2001

Reichardia tingitana* False sowthistle *

Senecio lautus Variable groundsel * Fg ‘3

Sonchus oleraceus* Common sowthistle * * *

Urospermum picroides* False hawkbit *

JUNCAGINACEAE

Triglochin mucronatum Prickly arrowgrass +

Triglochin tricophorum Arrowgrass %

LILIACEAE

Bulbine semibarbata Leek lily * * *

Dianella revoluta Black-anther flax lily * - *

var. revoluta

JUNCACEAE

Juncus bufonius Toad Rush +

GRAMINAE

Austrodanthonia White top *

caespitosa

Austrostipa flavescens Coast spear-grass *

Avena barbata* Bearded oat bu =

Bromus catharticus* Prairie grass *

Bromus diandrus* Great brome * ‘i

Cynodon dactylon* Couch-grass *

Digitaria violascens* Emu-grass * if

Elymus multiflorus Wheatgrass s

Hainardia cylindrica* Common barbgrass * *

Hordeum leporinum* Barley grass * * - *

Lachnagrostis filiformis Blown grass *

Lagurus ovatus * Hare's tail grass *

Lolium perennex Ryegrass =

rigidum*

Lolium perenne* Perennial ryegrass fe a

Lolium rigidum* Annual ryegrass r «

Parapholis incurva* Curly ryegrass 7 *

Poa fax Scaly poa i

Poa poiformis Coast tussock grass * if

var. poiformis

* oe

Sporobolus virginicus

Salt-couch

106 E. F LAWLEY & S. A. SHEPHERD

Methods

Surveys, photopoints and vegetation maps

In 2001 the survey of Althorpe I. described in this

paper was carried out using the methods of the

Biological Surveys of South Australia (Heard &

Channon 1997). The survey was allotted the

identifying number BS-126, in the Biological

Survey Data Base, where the collected data are now

stored as part of the Environmental Data Base of

South Australia.

Survey BS-126 comprises Althorpe I. (surveyed

June and October 2001) and the South Neptune Is

(surveyed October 2001). Numbered voucher

specimens of all species collected were identified and

deposited at the State Herbarium of South Australia.

Sites were chosen to represent the different vegetation

communities on the island, and quadrats located after

consulting aerial photographs and the previous floristic

map (Robinson ef al. 1996), and visual assessment. In

each of 10 standard 30x30 m quadrats on Althorpe I.,

all plant species were recorded, the dominant

overstorey and understorey species were noted, as well

as the life form and average plant height of each

species, using the adapted Muir’s table. Species’ cover

abundances were estimated using the modified Braun-

Blanquet scale (Heard & Channon 1997) and life

stages noted. The overall vegetation association in each

quadrat was described by summarizing the canopy

cover for the dominant strata using the codes from

Muir’s table, and the vegetation structure was described

from the SA Vegetation Structural Formations table

(Heard & Channon 1997). Names of vouchered

samples of plant species found outside the quadrats,

were added to the species list (Table 1).

Physical data including landform element, site

slope and aspect, outcrop cover, outcrop lithology,

surface strew-size, -cover and -lithology, fire-scars,

bare earth and plant litter coverage, and erosion were

also collected. Vertebrate presence and surface soil

texture class were examined and recorded, as was

disturbance within a 30 m radius of the quadrat.

The ten quadrats surveyed on Althorpe I. are shown

on the floristic map (Fig. 1). Eight of the 10 quadrats

are permanently marked with steel droppers carrying

the standard Biological Survey aluminium disks (Nos

9431-9438), as photopoints for future reference.

A 1999 aerial photo of Althorpe I. was ortho-

rectified using GPS coordinates of notable island

features, and used as a base to draw the vegetation

map. Boundaries between vegetation communities

were ground-truthed using handheld GPS.

Results

Vegetation Mapping

The vegetation map (Fig. 1) presents five distinct

native vegetation formations and a non-native grass

community, which are described below.

Nitre bush (Nitraria billardierei) Low SHRUBLAND

(sites ALT00301, ALTO0401, ALTO0501) — Fig. 2A.

This occurs on the central and eastern area of the

main plateau of Althorpe I. that is covered in deep

sand with occasional low dunes. Seaberry saltbush

(Rhagodia candolleana ssp. candolleana), and coast

twin leaf (Zygophyllum billardierei), occur in almost

equal cover abundance as the nitre bush (N.

billardierei), which is the tallest naturally occurring

native species on the Island. Groundcovers are the

native perennials, coast bone fruit (Threlkeldia

diffusa), and ruby saltbush (Enchylaena tomentosa

ssp. tomentosa) as well as some introduced annual

forbs and grasses.

African boxthorn (Lycium ferocissimum) was well

established here until 1998, when it was

progressively removed. The area is burrowed by

short-tailed shearwaters at an average density of 12-

15 burrows per 100 m?.

Low OPEN

ALT 00201,

Atriplex paludosa

SHRUBLAND _ (sites

ALT01001) — Fig. 2B.

This chenopod shrubland occurs in shallow soils

over calcarenite. Rhagodia candolleana ssp.

candolleana, an associated plant, increases in

percentage of cover where the soil deepens, whilst

conversely the mat plant Disphyma crassifolium ssp.

clavellatum decreases. Species on the island unique

to this shrubland are the native forbs, hairy plantain

(Plantago hispida), billy-buttons (Craspedia

variabilis), and yam daisy, (Microseris lanceolata).

Some native perennial grasses and introduced annual

grasses and medics also occur in this Atriplex

shrubland. Short-tailed shearwater burrows occur in

this habitat at average densities of 6-8 per 100 m?.

ssp. cordata

ALT00101,

Maireana oppositifolia and Atriplex paludosa ssp.

cordata LOW SHRUBLAND (Sites ALTOO801 and

ALT00901) — Fig. 2C.

This low shrubland is on the island’s very exposed

steep slopes, but extends onto the plateau in places

where the surface calcarenite is exposed or covered

by a thin veneer of soil. Robinson ef al. (1996)

classified this shrubland as Leucophyta brownii

Cushion bush (sub nom. Calocephalus brownii), but

this species is quite uncommon and absent from

many areas on the slopes. Its cover abundance was

classified in quadrat ALTOO801 as N (1-10

individual plants) and in ALTO0901 as zero.

Maireana oppositifolia and Atriplex paludosa ssp.

cordata had a cover of 5 — 25% in quadrat

ALT00801, and 25 — 50% in ALTO0901.

In sheltered pockets in the calcarenite surface

VEGETATION OF THE ALTHORPE ISLANDS 107

many other ground cover and shrub species occur,

such as round-leaf pigface (Disphyma crassifolium

ssp. clavellatum), coast bone fruit (Threlkeldia

diffusa), seaberry saltbush (Rhagodia candolleana

ssp. candolleana), coast twin leaf (Zygophyllum

billardierei), bassia (Sclerolaena uniflora), southern

sea heath (Frankenia pauciflora var. fruticulosa) and

coast plover-daisy (Leiocarpa supina), in addition to

grasses and numerous herbaceous plants. Less

frequent on the slopes are the vivid purple flowering

coast swainson-pea (Swainsona lessertiifolia), and

the large golden flowering pleated podolepis

(Podolepis rugata var. littoralis). The coastal Lrodia

achillaeoides ssp. achillaeoides has been found only

on the northern slope west of the saddle. The quarry

survey site ALT 00701 (Fig. 1) is now re-vegetated

with this low Maireana/Atriplex shrubland. N.

billardierei grows densely and in substantial patches

on the slopes where rubble fall has increased soil

depth, but is at too small a scale to map. The

Maireana/Atriplex. community grades into the

Atriplex paludosa ssp. cordata vegetation type where

the soil deepens.

Halosarcia pergranulata Low CLOSED SHRUBLAND

This vegetation formation, absent from Robinson

et al. (1996), is only found near sea-level on the

northern point, called ‘The Monuments’, where

calcarenite outcrops over a granitic base partly

shelter the site from the prevailing SW _ winds.

Species present are: samphire (Halosarcia

pergranulata), with some groundcover of sea celery

(Apium prostratum var. filiforme), Frankenia

pauciflora var. fruticulosa, Enchylaena tomentosa

var. tomentosa, and the common _ iceplant

(Mesembryanthemum — crystallinum). —Karkalla

(Carpobrotus rossii) grows thickly out onto granite

edging this site.

Myoporum insulare LOW OPEN SHRUBLAND (site

ALT00601) — Fig. 2D.

This formation, absent from Robinson ef al.

(1996), is located on sand at the isthmus to ‘The

Monuments’. Sand over granite, grading to almost

bare granite, supports 24 species of which 8 are

introduced. Predominant are common boobialla

(Myoporum insulare), Nitraria billardierei, coastal

climbing lignum (Muehlenbeckia gunnii), Atriplex

paludosa ssp. cordata and Leiocarpa supina, with

groundcovers Carpobrotus rossii and Disphyma

crassifolium ssp. clavellatum. Sparsely represented

are: sea spurge (Euphorbia paralias), two-horned sea

rocket (Cakile maritima), Mesembrvanthemum

crystallinum, and four other herbaceous non-natives.

® Census of South Australian Vascular Plants. Ed 5.1. Staff and

Assoc. of the State Herb. of South Australia, May 2004.

A few grey saltbush (Atriplex cinerea), the only

specimens on the Island, and some Australian

hollyhock (Malva behriana), were also found here.

The once present African boxthorn (Lycium

ferocissimum), has been eradicated.

Non-native Grasslands and Cleared Areas

This cover type encompasses the landing strip and

the area taken up by buildings and yards (Fig. 1). The

vegetated area, of mainly annual non-native grasses,

is found in a patch on the eastern cliff top and also on

the landing strip, where in addition to the annuals,

Cynodon dactylon, a perennial non-native couch

grass, occurs. Maintenance of the landing strip by

mowing ceased in 2001, 10 years after the light-

station was automated, and shearwater burrows are

gradually reappearing in the area.

Island Perimeter

Perpendicular sandstone cliff faces and steep

calcarenite slopes, often covered in talus scree form

the island’s perimeter. A typical shoreline vegetation,

too narrow to be accurately mapped, covers the area

between the foot of the slopes and the granite

intertidal. Common species are: Cakile maritima,

Disphyma crassifolium ssp. clavellatum, Apium

prostratum var. filiforme, Carpobrotus rossii, and

where the soil is deeper, Nitraria billardierei and the

occasional Myoporum insulare or Malva behriana

shrub.

Species With Few Records

Peppercress (Lepidium foliosum), collected by

White (1916) on Althorpe and Seal Is, has not been

recorded since then. Stirling ef a/. (1970) recorded

the same species on the South Neptune Is, where it

was also absent in 2001. Keighery ef a/. (2002) noted

that Lepidium foliosum grew in association with

guano-rich seabird nesting sites. Small-leaved

mintbush (Prostanthera — serpyllifolia ssp.

serpyllifolia), recorded by Robinson ef al. (1996),

has not been found again, despite intensive searches.

The native grass, Elymus multiflorus, previously

recorded only in the Flinders Ranges (FR) and

Eastern (EA) areas of the State (Census 2004°), was

also recorded on Althorpe I. in 2001.

In all, 86 plant species are recorded from Althorpe

I., of which 46 are native and 40 introduced.

Discussion

Vegetation change over time, introductions, removals

and regeneration

The changes in Althorpe I.’s vegetation must have

been severe following human settlement. Goats

arguably had the greatest impact on the vegetation,

even that on the steep slopes, as witnessed by the

108 E. F LAWLEY & S. A. SHEPHERD

Fig. 2. Photos of four of the vegetation communities on Althorpe Island. A: Nitraria billardierei_ Low SHRUBLAND. B:

Atriplex paludosa ssp. cordata Low OPEN SHRUBLAND. C: Maireana oppositifolia and Atriplex paludosa ssp. cordata

Low SHRUBLAND. D: Myoporum insulare LOw OPEN SHRUBLAND.

many goat tracks still visible many years after goat

eradication. South Neptune I. has a_ similar

occupation history, and Dr. R. S. Rogers wrote in

1907, only six years after the establishment of its

Lighthouse: “There is a large number of big goats on

the island, and these animals have already changed

the vegetation” (Maiden 1908).

Besides the impact of goats and sheep, more subtle

changes may have been caused by the disappearance

(Medlin’) of the herbivorous greater-sticknest rat

(Leporillus conditor), and the arrival of garden snails

(Helix asper), house mice (Mus musculus), and

introduced plants on the island. Although the species

present prior to settlement in 1879 can never be

known, it may be inferred from a floristic

comparison with other islands and the mainland

coast, and from records of species’ responses to

grazing or competition from introduced species.

Adair & Groves (1998) found that most Australian

studies quantifying weed impact demonstrated a

decline in species richness, canopy cover or

frequency of native species.

Introduction of grazers may reduce native

vegetation and lead to extinctions of native plant

species, and the removal of the same grazers can then

lead to the spread of invasive exotics (Cox et al.1967

in Timmins & Geritzlehner 2003), and subsequent

loss of those native species vulnerable to

competition.

Abbott et al. (2000) found a 37% reduction in plant

species over 40 years on Carnac I., Western Australia

since 1951. They attributed the remarkable

vegetational changes inter alia to the removal of

rabbits and the arrival of the introduced species,

Mesembryanthemum crystallinum, Malva parviflora,

and more recently, Lycium ferocissimum. Althorpe I.

is likely to have suffered a similar loss of native plant

species.

Comparison with the mainland coast shows that

Althorpe I. has all but one, Melaleuca lanceolata, of

the eight perennial plant species, which occur in all

geomorphic regions along the coast (Opperman

1999). Species likely to have been on Althorpe I. at

the time of human settlement include some of the 12

species that occurred in over 25% of the SA Coastal

survey quadrats (Opperman 1999). These are:

Dianella brevicaulis, Olearia axillaris, Leucopogon

parviflorus, Isolepis nodosa, Pimelea serpyllifolia

ssp. serpyllifolia and Clematis microphylla. The

island’s physical environment and soils are also

suitable for growth of Exocarpos aphyllus, Beyeria

lechenaultii, Scaevola crassifolia, Helichrysum

VEGETATION OF THE ALTHORPE ISLANDS 109

leucopsideum, Alyxia buxifolia and, at the base of the

island, Swaeda australis. The disappearance of

Lepidium foliosum might be due to the 1980s

decline of the fairy penguin (Eudyptula minor), and

the consequent loss of bird guano favoured by this

species (Keighery ef al. 2002).

The South Neptune Islands (SNI) species list

(Table 1) provides further insight. Seven SNI North

I. native species were not found, and may have

disappeared from SNI Lighthouse I.'°, the most

obvious being Correa reflexa var. reflexa and coastal

lignum, Muehlenbeckia gunnii. The latter species has

persisted on Althorpe I. in very low numbers, notably

in crevices among rocks. On SNI North I. coast

tussock grass, Poa poiformis, forms an extensive

native grassland community, but it is rare on SNI

Lighthouse I., possibly due to heavy grazing by

resident Cape Barren Geese preventing its re-

establishment after goat removal. These birds rarely

visit Althorpe I. and do not breed there. P poiformis,

which requires a certain level of salt spray

(Opperman 1999), is absent from Althorpe I. despite

the presence of suitable habitats on its lower slopes.

It was possibly grazed to extinction. A comparison

with the previously occupied 809 ha St Francis I.,

Nuyts Archipelago, shows some habitats similar to

those on Althorpe I. (Robinson e¢ al. 2003). The

Nitraria billardierei community on the cliff top

sands on St Francis I. contains Olearia axillaris and

Myoporum insulare, both missing from the

equivalent community on Althorpe I. Of the 131

plant species recorded from St Francis I. 26% are

exotic (Robinson ef al. 2003) compared with the

47% of exotic species among the 86 species on

Althorpe I. (Table 1). This difference may be due to

the greater proximity of Althorpe I. to the mainland,

its smaller size or greater modification.

The spatial extent of some species has apparently

also become much reduced. Bower spinach, 7etragonia

implexicoma, is usually a common and widespread

coastal and island species, which may rapidly go

extinct under grazing (Underwood & Bunce 2004), It

has survived on Althorpe I. only on the moister, steep,

southerly-facing slopes intensely burrowed by

shearwaters, and unattractive to grazing goats.

Two native grasses are poorly represented on

Althorpe I. Coast spear-grass, Austrostipa flavescens,

occurring in various areas on shallow sand near the

plateau perimeter, is favoured by the shearwaters as

nesting habitat, possibly because the tussocks bind

the soil, preventing erosion and burrow collapse. The

palatable and nutritious white-top Austrodanthonia

caespitosa, (see Jessop & Toelken 1986) is now

‘© A number of native species are recorded for SNI Lighthouse I. but

not recorded for SNI North I., they are possibly present but survey

time on North Island was 8 hours, on Lighthouse Island 10 days.

restricted to a few sites on the cliff top edges, and

was likely grazed to near extinction. Exotic annual

grasses are now widespread on Althorpe I. and may

bind the soil and reduce soil erosion in areas used by

shearwaters.

Exotic grasses were also sown on the newly graded

landing strip at South Neptune Lighthouse I. in 1961,

but failed to establish there and persist only among

granite boulders at the northern end of the strip.

Since goat eradication by 1990 on Althorpe I. an

increase in cover of native species has been

observed. Whereas Robinson ef al. (1996) noted that

introduced grassland at the western end of the

plateau was common in 1982, by 2001 Atriplex

paludosa ssp. cordata, had become the predominant

canopy species (Fig. 1). Nitraria billardierei had also

spread, obliterating former tractor tracks, likely

because the large seed in its palatable fruit is readily

spread by gulls and other birds (unpublished

observations). Stirling et al. (1970) also observed N.

billardierei seed in regurgitate of the Pacific gull,

Larus pacificus, on North South Neptune I. between

1967-1970, although N. billardierei was then absent.

However, by 2001 the species had established there

in small numbers.

The old landing strip on Althorpe I. now has the

groundcovers, Threlkeldia diffusa and Enchylaena

tomentosa var. tomentosa among the perennial couch

grass, (Cynodon dactylon), and annual grasses. An

African boxthorn control program has been in

operation since 1998 and the introduced herbaceous

biennial, tree mallow, Malva dendromorpha, once

common on the plateau, has been manually removed

since 1996, and is now in low numbers.

Conclusions

The vegetation of Althorpe I. has been severely

disturbed by goats and human occupation between

1879 and 1991, with the resultant likely loss of native

species and the introduction of many exotics. Since

1991, greater environmental awareness has

stimulated efforts to eliminate introduced animals,

and eradicate exotic perennials. Goat eradication and

boxthorn control on Althorpe I. are beginning to

show benefits, and providing that boxthorn re-

establishment is prevented, the documented trend of

native vegetation recovery is expected to continue.

The 2001 and later vegetation surveys, with fixed

photopoints, have established a valuable data-base

and baseline, against which to compare future

changes in the vegetation.

Acknowledgements

We thank the Native Vegetation Council for the

grant enabling the 2001 vegetation survey, Innes

110 E. F LAWLEY & S. A. SHEPHERD

National Park staff for cooperation and support, Lee

Heard and Sue Kenny of (then) Planning SA, for

training Friends of Althorpe I. Conservation Park,

supporting the 2001 survey and entering electronic

data. Rosemary Taplin and Helen Vonow at the State

Herbarium of South Australia assisted with

identifications, searched early records, and resolved

many difficulties. Sue Kenny helpfully modified

Figure 1, Peter Canty provided information and

advice on the 1982 survey, John Lawley provided

historical information, Sherilee Hawkins carried out

preliminary survey work, and Brad Page provided

much helpful advice in assembling the data. We also

thank Rob Lewis and Anthony Cheshire of SARDI

for promoting the Island research, and Sue Murray-

Jones for her administrative skills. Wendy Stubbs,

Helen Vonow, Bob Baldock and referees provided

helpful comments on the manuscript.

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term change in the floristic composition and vegetation

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Environmental Weeds on Biodiversity, A Review and

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Bevperio, A. P., HAILS, J. R. & Gostin, V. A. (1983) A

review of Holocene sea levels in South Australia. /n

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Australia Methodology. First Edition”. (Geographic

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Australia” 4th ed. South Australian Government Printer

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KEIGHERY, G. J., ALFORD, J. J. & LONGMAN, V. (2002) A

vegetation survey of the islands of the Turquoise Coast

from Dongara to Lancelin, south-western Australia.

Conservation Science W. Aust. 4, 13-62.

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Island Biogeography”. (Princeton University Press,

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Australia. Trans. R. Soc. S. Aust. 32, 252-286.

OPPERMAN, A. (1999) “A Biological Survey of the South

Australian Coastal Dune and Clifftop Vegetation” (Coast

and Marine Section, Environment Protection Agency,

Adelaide).

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“South Australia’s Offshore Islands”. Australian Heritage

Commission. (Australian Government Publishing

Service, Canberra).

RoBINSON, A. C., CANTY, P. D., WACE, N. M. & BARKER R.

M. (2003) The Encounter 2002 expedition to the Isles of

St Francis, South Australia: Flora and vegetation. Trans.

R. Soc. S. Aust. 127, 107-128.

STIRLING, I., STIRLING, S. M., & SHAUGHNESSY, G. (1970)

The bird fauna of South Neptune Islands, South

Australia. Emu 70, 189-192.

TIMMINS, S. & GERITZLEHNER, J. (2003) Do weeds respond

to Pest Animal Control? DOC Science Internal Series

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Unpberwoop, M. & BuNCcE, A. (2004) The breeding biology

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Transactions of the Royal Society of S. Aust. (2005), 129(2), 111-115.

EFFECTS OF AFRICAN BOXTHORN REMOVAL ON NATIVE VEGETATION

AND BURROWING OF SHORT-TAILED SHEARWATERS ON ALTHORPE ISLAND,

SOUTH AUSTRALIA

by E. FE. Lawtey!, J. J. LAWLEY? & B. PAGs

Summary

Law ey, E. F, LAwLey, J. J. & Pace, B. (2005). Effects of African boxthorn removal on native vegetation and

burrowing of short-tailed shearwaters on Althorpe Island, South Australia, Trans. R. Soc. S. Aust. 129(2), 111-115,

30 November, 2005.

Althorpe Island’s African boxthorn infestation was incrementally removed from 1998 and the effect of this

removal on the vegetation and on the nesting of the short-tailed shearwater, Puffinus tenuirostris, was examined.

During the study period, native perennial mat plant vegetation succeeded an initial flush of annual non-native

plants at most sites examined. A weak trend of increasing burrow density of shearwaters within two metres from

boxthorn stumps after removal of the canopy was also observed.

Key Worps: Lycium ferocissimum; African boxthorn; Puffinus tenuirostris; short-tailed shearwater; succession;

Althorpe Island

Introduction

The increased soil nutrient loads (through guano)

and disturbance (excavating and trampling) caused

by seabirds have been shown to decrease both the

density of plants and the species richness in seabird

colonies (Crooks 2002; Bancroft et al. 2005).

Vegetation within colonies of burrowing seabirds is

typically dominated by only a few species (reviewed

in Walsh ef al. 1997). Within Australian colonies of

shearwaters (Puffinus spp.) the dominant vegetation

is typically the succulents: Tetragonia spp., Rhagodia

spp. and/or Carpobrotus spp. (Walsh et al. 1997).

However, where soil nutrient loads are naturally low,

the combination of increased guano deposition and

disturbance are thought to create conditions that

favour annual and/or exotic plant species (Lamont

1984). In that study, a shearwater colony in Western

Australia, was dominated by short-lived, succulent

exotics, whereas areas surrounding the colony were

dominated by long-lived heath species.

The removal of invasive alien species from natural

ecosystems can have unexpected outcomes. If the

alien species are at different trophic levels, then

mesopredator release, cascade and other indirect

effects can occur in the system after their removal

(Zavaleta et al. 2001). Hence it is critically important

to undertake post-eradication monitoring in order to

assess the effect of eradication.

Althorpe I. (Lat. 35° 23’ S; Long. 136° 51’ E),

located 8.5 km off Cape Spencer on Yorke Peninsula,

' PO Box 89, Tarlee, South Australia 5411.

Email: lawley@chariot.net.au

> Tbid

} SARDI Aquatic Sciences, PO Box 120, Henley Beach, South

Australia 5022

South Australia, is 92 ha in area. The island has high

conservation value as it is the nesting site of about

20,000 — short-tailed = shearwaters, — Puffinus

tenuirostris, which dig nesting burrows in the sandy

soil (Robinson ef al. 1996). Cats, Felis catus, and

goats, Capra hircus, were introduced during human

occupation (Lawley & Shepherd 2005) and may have

affected shearwater breeding success by predation

and trampling respectively. The goats were

eradicated by 1990, and the cats by 2003.

The African boxthorn, Lycium ferocissimum,

introduced to Althorpe I. around the 1930s,

proliferated after the removal of goats, and became

abundant in virtually all habitats (Lawley &

Shepherd 2005). Observations suggested that

shearwaters were unable to burrow under the

boxthorns, which develop a fine, dense root mass

under the bole. From 1998 a boxthorn eradication

program was undertaken. An estimated 10000

mature boxthorns were removed, and in the

succeeding six years up to 2000 seedlings were

removed annually.

Following the boxthorn eradication program, this

study was undertaken to determine:

(1) the effect of boxthorn removal on_ the

surrounding vegetation, and in particular, the

process of recolonisation in the bare patches

produced by boxthorn removal; and

(2) whether shearwaters expanded their burrowing

activities into the above bare patches.

Materials and Methods

Recolonisation of bare patches

Twenty stumps of large boxthorns, left in situ on

removal of the canopies, were selected for

112 E. EF LAWLEY, J. J. LAWLEY & B. PAGE

monitoring in May 2001. Each stump was marked

with paint and a tag bearing the year of clearing and

a sequential number, and its GPS location recorded.

A hoop of 1.5 m diameter, divided into segments to

facilitate estimation of cover, was placed around the

stump, and the enclosed quadrat photographed. The

number of plants of each species appearing in the

quadrats was recorded in three categories: recently

germinated seedlings; immature plants >5 cm high;

and mature plants. For spreading perennials, the

percent cover was estimated visually. Percentage

cover of annual seedlings was not plotted because

germination date and growth rate of these are closely

linked to autumn rain events, the onset of which may

vary by several weeks. A difference in cover of

annuals at the time of the survey may not be a true

reflection of the temporal trend. The quadrats were

resampled in the spring of 2001, and in the autumn

and spring of 2002 and 2004.

Shearwater burrow density

Studies were concentrated solely in a Nitraria

billardierei shrubland where burrows were deepest,

to about | m depth, and at relatively high densities.

Four transects (named A-D), each 60 m long (except

transect B which was 45 m), were set on level

ground, with similar densities of boxthorn among

uniform native vegetation, and with similar soil

depths. Transect D was placed in an area cleared of

boxthorn in 2000, A in an area cleared in 2001 and

Transects B and C in areas cleared in early 2002.

GPS coordinates for the southern extremities of

transects were noted, and marked with permanent

droppers with tags recording the transect number and

year of clearing. Distances between shearwater

burrows and boxthorn stumps were measured in

April 2002, 2003 and 2004 according to the

following protocol. Observers walked from south to

north on either side of a tape measure stretched

between the extremities of the transect, holding a 2.5

m measuring stick at right angles to the tape.

The exact location of each boxthorn stump or bush

with a trunk diameter >50 mm, within 2.5 m of the

transect line, was recorded. Shearwater burrows were

included in the study if they showed signs of having

recently been used by shearwaters (guano deposits,

recent excavations and/or chick present). Each

shearwater burrow’s distance to a stump, within a

radius of 2 m of each stump, was recorded by

measuring from the centre of the burrow entrance to

the centre of the nearest stump, to an accuracy of

10 cm, in the categories 0 — 0.5 m, >0.5 — | m, or

>1— 2 m distance from stumps. In Transects A-D

there were 6, 8, 14 and 10 stumps respectively. Based

on the methods outlined above, the mean burrow

density was calculated for each transect, in each of

the distance categories from stumps (0 — 0.5 m =

0.79 m’, >0.5 — 1 m = 2.36 m2, or >1 — 2 m= 9.43

m?). The number of burrows around boxthorn stumps

was recorded from the period before boxthorn

removal until four years after removal (Fig. 1).

To monitor average burrow density over time for

each transect area, a burrow count was conducted in

a 10 x 10 m quadrat at the ends of each transect.

Statistical analyses, including the Spearman rank

correlation, r, were done with the SPSS statistical

software (version 11, SPSS Inc., Chicago). All

statistical tests are two-tailed, unless stated, with the

o level of statistical significance set at 0.05.

Results

Spontaneous recolonisation

In the first year after boxthorn removal, many

species appeared in the quadrats, including annual

non-native forb and grass seedlings, such as common

sowthistle, Sonchus olearaceus (up to 260 seedlings

per quadrat), ryegrass, Lolium sp. (up to 120

seedlings per quadrat), common __ iceplant,

Mesembryanthemum crystallinum (up to 200

seedlings per quadrat). Perennial native seedlings

persisted through the summer, and, together with

vegetative expansion of plants outside the quadrats,

an overall increase of cover by perennial native

plants was observed over the four years observation

period (Fig. 2 and Fig. 3).

Plots showing the changes in mean percent cover

of the four most common native species over four

years are given in Fig. 3. Seaberry saltbush,

Rhagodia candolleana ssp. candolleana (Fig. 3A),

appeared in six quadrats, showing an increase in

percent cover in five quadrats, but disappeared from

the sixth where it was overgrown by the common

iceplant, Mesembryanthemum crystallinum, and wild

turnip Brassica tournefortii. A mat plant, coast bone

fruit, Threlkeldia diffusa (Fig. 3B), colonized rapidly,

appearing in nine quadrats after one year, and in 17

after four years. Another mat plant, ruby saltbush,

Enchylaena tomentosa var. tomentosa (Fig. 3C),

appeared in one quadrat after one year, but was

present in 11 quadrats after four years. Coast

twinleaf, Zygophyllum billardierei (Fig. 3D), initially

appeared in seven sites but subsequently declined in

some quadrats. By spring 2004 some of the stumps,

four years after cutting, had disappeared completely

under the native matplants with a cumulative cover

of almost 100% (Fig. 2).

Shearwater burrow density

In the N. billardierei shrubland overall burrow

density, as measured in the quadrats at the ends of the

transects, ranged from 0.12 — 0.19 burrows m*°.

There were too few data to examine inter-annual

changes in burrow density. Along the transects, the

BOXTHORN REMOVAL EFFECTS ON NATIVE VEGETATION AND SHEARWATERS 113

0.30

0.25

0.15

Number of burrows per square metre

0.10

0.05

—~o—0.0 to 0.5m

tA 05to1.0m

—B—1.0to2.0m

—B— 0.0 to 2.0m

a er eee |

1 year prior 1 year after 2 years after 3 years after 4 years after

[ Number of years since boxthorn removal

Fig. |. The mean number of short-tailed shearwater burrows around individually numbered boxthorn stumps, measured

before and after boxthorn removal. For clarity, standard error bars are shown only for summed density data, 0 — 2 m from

stumps.

mean burrow density was positively correlated with

the number of years since boxthorn removal;

however, none of the regressions were significant

(0-0.5 m: r = 0.286, df = 3, P = 0.36; 0.5 — 1.0 m:

r= 0.417, df = 3, P = 0.29; 1.0 — 2.0 m: r = 0.686,

df = 3, P = 0.16; Fig. 1). The overall mean burrow

density between 0 — 2 m was positively correlated

with the number of years since boxthorn removal and

the regression approached significance (0 — 2.0 m:

r= 0.833, P = 0.08, Fig. 1).

Discussion

The eradication of goats from Althorpe I. most

likely had pervasive ecosystem consequences before

this study began. Boxthorn plants, previously kept at

low levels by goats, proliferated after their

eradication, and boxthorn became abundant in

virtually all habitats. The rapid spread of boxthorn

was aided by at least two vectors common on

Althorpe L., the little raven, Corvus mellori, and the

starling, Sturnus vulgaris, which eat boxthorn berries

and spread seed in regurgitates and droppings (EFL,

unpublished data).

In this study, the immediate effect of boxthorn

removal was the exposure of bare patches. Apparently

at Althorpe I., the shading by the boxthorn canopy,

combined with its extensive root system, was able to

out-compete and eliminate understorey growth (see

Belsky 1994, Holmgren et al. 1997). After clearing of

boxthorns, annual weeds invaded the clearings, and in

succeeding years were gradually replaced by native

perennials in most of the 20 quadrats (Fig. 3). A

similar succession of exotic winter annuals, followed

by native mat plants in bare areas, was found by

Brown ef al. (1993) in Tasmania. However, in that

study the bare areas were a consequence of

shearwater trampling, not boxthorn removal.

114

E. RF LAWLEY, J. J. LAWLEY & B. PAGE

10 Rhagodia candolleana

fay

fo)

0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 45

15 Enchylaena tomentosa

o@ 10

fe}

is)

0 05 1.0 1.5 2.0 2.5 3.0 3.5 4.0 45

Number of years since boxthorn removal

Threlkeldia diffusa

1p Zygophyllum billardierei

0) r — r 1 1

0 05 1.0 1.5 20 2.5 3.0 3.5 4.0 45

Number of years since boxthorn removal

Fig. 3. Mean percent cover of Rhagodia candolleana (A), Threlkeldia diffusa (B), Enchylaena tomentosa (C), and

Zygophyllum billardierei (D) in marked quadrats, following the removal of boxthorn (Lycium ferocissimum). Vertical bars

are standard errors.

Whilst this study has shown only a weak temporal

trend in increased burrowing near boxthorn stumps,

this may be because too few years have elapsed for

the fibrous root system under the stumps to break

down sufficiently for shearwaters to burrow through

them. Further, more time may be needed for young

shearwaters to excavate new burrows in the greater

area made available.

The shearwater burrow densities in the quadrats at

the end of each transect encompassed the range

observed in the study sites, particularly those

observed up till two years after removal. The data, as

yet inconclusive, suggest that burrow density is

rising.

BOXTHORN REMOVAL EFFECTS ON NATIVE VEGETATION AND SHEARWATERS 115

Conclusions

The removal of African boxthorn has had a positive

effect on the native flora, by allowing native shrubs,

particularly mat plants, to recolonise the bare

patches. Whether shearwaters have expanded their

burrowing activities into the areas vacated by

boxthorn is less clear. Continued monitoring,

perhaps triennially, both of shearwater burrow

density and of the plant quadrats, is recommended to

determine long-term trends.

Short-tailed shearwater colonies are absent from

the southern Australian mainland and found only on

a few suitable offshore islands. The destruction or

degeneration of habitat can cause the demise of these

sea-bird colonies, and Jones ef al. (2003)

documented such declines in New Zealand, largely

due to introduced predators. Hence, the eradication

of feral cats and the unremitting control of

aggressive alien species, such as boxthorn, will help

conserve both the native vegetation and seabird

colonies on these islands.

Acknowledgements

We thank the Friends of Althorpe Islands

Conservation Park for facilitating the field trips and

participation in fieldwork, and the staff at Innes

National Park for support. We also thank Rob Lewis

and Anthony Cheshire of SARDI for promoting the

Island research, Scoresby Shepherd for

encouragement, helpful comments and for editing

the manuscript, and referees for improvements to the

paper.

References

Bancrort, W. J., ROBERTS, J. D. & GARKAKLIS, M. J. (2005)

Burrowing seabirds drive decreased diversity and

structural complexity, and increased productivity in

insular-vegetation communities. Aust. J. Botany 53, 231-

241.

Betsky, A. J. (1994) Influences of trees on savanna

productivity: tests of shade, nutrients, and tree-grass

competition. Ecology 75, 922-932.

Brown, M. J., MARUYAMA, N. & WILLIAMS, K. J. (1993)

Ecological studies of vegetation in short-tailed

shearwater colonies in Tasmania. Pap. Proc. R. Soc.

Tasm. 127, 11-16.

Crooks, J. A. (2002) Characterizing ecosystem-level

consequences of biological invasions: the role of

ecosystem engineers. Oikos. 97, 153-166.

Lamont, B. B. (1984) Specialised modes of nutrition. pp.

126-145 In Pate, J. S. & Beard, J. S. (Eds) “Kwongan,

Plant Life of the Sandplain”. (University of Western

Australia Press, Perth).

HOLMGREN, M., SCHEFFER, M. & Huston, M. A. (1997) The

interplay of facilitation and competition in plant

communities. Ecology 78, 1966-1975.

Jones, C., BETTANY, S., MOLLER, H., FLETCHER, D. & Cruz,

J. D, (2003) Burrow occupancy and productivity at

coastal Sooty shearwater (Puffinus griseus) breeding

colonies, South Island, New Zealand: can mark-

recapture be used to estimate burrowscope accuracy?

Wildlife Research 30(4), 377-388.

Lawey, E. F & SuHepHerp, S. A. (2005) Land use and

vegetation of Althorpe Island, South Australia, and a

floristic comparison with South Neptune Islands. Trans

R. Soc, S. Aust. 129, 100-110.

Rosinson, A. C., CANTY, P., Mooney, P. & Rupbuck, P.

(1996) “South Australia’s Offshore Islands”. Australian

Heritage Commission (Australian Government

Publishing Service, Canberra).

ZAVALETA, E. S., Hops, R. J. & Mooney, H. A. (2001)

Viewing invasive species removal in a whole-ecosystem

context. Trends in Ecology and Evolution 16, 454-459.

Watsh, J. B., Kirkpatrick, J. B. & Skira, I. J. (1997)

Vegetation patterns, environmental correlates and

vegetation change in a Puffinus tenuirostris breeding

colony at Cape Queen Elizabeth, Tasmania. Aust. J.

Botany 45, 71-79.

Transactions of the Royal Society of S. Aust. (2005), 129(2), 116-127.

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

by R. N. BALDock! & H. B. S. WOMERSLEY?

Summary

Batpock, R. N, & WomersLey, H. B. S. (2005) Marine benthic algae of the Althorpe Islands, South Australia.

Trans, R. Soc. S. Aust. 129(2), 116-127, 30 November, 2005.

Deep water to intertidal marine algae have been collected from the Althorpe Islands on four occasions. Present

collections include 15 species of Chlorophyta, 37 Phaeophyta and 92 Rhodophyta, probably a fraction of the

total species present. A canopy of Acrocarpia paniculata and Ecklonia radiata occurs in most subtidal regions.

Algal communities at a cave site and those exposed to high wave energy and sheltered conditions are described.

Kry Worpbs: Marine benthic algae; Althorpe Islands; offshore islands; algal communities; intertidal algae;

subtidal algae; algal canopy

Introduction

The Althorpe Islands lie in Investigator Strait,

South Australia and consist of a single, large, steep-

sided calcarenite island and several lower outlying

isles. The group was separated from southern Yorke

Peninsula, S. Australia, by sea level rise during the

last interglacial period. They are underlain by jointed

granites and amphibolite dykes (see Robinson ef al.

1996, p.288) supporting vertical, orange, calcarenite

cliffs with caves on exposed coastlines. Unlike parts

of the adjacent mainland there are no flat calcarenite

shore platforms, and only a single relatively narrow

sandy beach, the Mooring Bay. A granite rise with

joints and dykes on the western side has been eroded

to five isles, the Western Isles. Subject to the full

effects of cyclonic cells moving east, the Althorpes

are exposed to periodic high wave energy on all

sides.

Algae deposited in the State Herbarium have also

been collected from the adjacent southern Yorke

Peninsula mainland over a number of years by

Shepherd & Womersley, and Edyvane & Baker

collected at nearby Haystack, Chinaman Hat and

Seal Islands for the construction of the Interim

Marine and Coastal Regionalisation of Australia

(IMCRA). (See Edyvane & Baker 1996.)

An Investigator Strait collection (J. Watson,

January 1971, unpublished) from two deep-water

transects between Foul Bay/Hillock Point on the

mainland and Cape Cassini/Cape Dutton on

Kangaroo Island has also been made.

The present study although confined to the algae of

the main Althorpe Island and its five outliers, the

' University of South Australia, Mawson Lakes campus, South

Australia, 5095. Email: Robert.Baldock@unisa.edu.au, and State

Herbarium.

2 State Herbarium, Botanic Garden, Hackney Rd, Hackney, South

Australia 5000, and Department of Environmental Biology,

University of Adelaide.

Western Isles, brings together data from both the

2004 expedition and several previous studies to

describe algal distribution of these islands.

Collections for these studies have been made by

the following:

1. By divers aboard the vessel SAORI, 4 January,

1964 while undertaking a marine exploration of

S. Australian Gulf waters;

2. By J. Baker and K. Edyvane, 26-27 October,

1993, during the collection of data for IMCRA;

3. By A. Gaut, 2001 from the intertidal, but also

containing drift specimens;

4. By E. Lawley, 28 September, 2001 and 28

November, 2004 from a saline soak 50 m inland

and 10 m above sea level;

5. By R. Baldock, J. Brook, R. Lewis and

N. Barrett, January 31 — February 11, 2004,

mainly using SCUBA during the expedition

conducted by the South Australian Research and

Development Institute (SARDI) aboard the

research vessel Ngerin.

There is also a single specimen of Cystophora

platylobium collected by M. Retallick, from 18 m

deep off the N. end, on 3 March 1987.

Collation of collections in the current study will

permit a database for further ecological work and

comparisons with other local regions such as the

Gambier Island group, suggested for Marine

Protected Areas.

Names and classifications below follow “The

Marine Benthic Flora of Southern Australia”

(Womersley 1984, 1987, 1994. 1996, 1998, 2003),

with changes from later phycological studies.

Methods

In 2004, seven collecting sites were chosen on the

basis of accessibility and contrasting habitat

conditions. They have been described in terms of

physical conditions and floristics; 2004

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 117

identifications of species are generally by the first

author, and the remainder by H. B. S.Womersley,

who checked all determinations. The species list

consists exclusively of specimens deposited in the

State Herbarium.

Biomass differences at several depths between

“The Boulders”, with moderate wave energy, and

“The Hump”, a high energy habitat, were also

determined in order to expand the site descriptions.

Sub-tidal algae were collected from four 1 m?

quadrats at several depths with the help of A. Hirst,

then identified and weighed. Encrusting lithophytic

algae and sea grasses were not collected.

Results

Site Descriptions for the 2004 Collections

I. “The Boulders” (BO), E corner of Mooring Bay

This is a granitic headland exposed to moderate

wave energy and tidal current. The substratum slopes

at about 10° to a sandy bottom at about 8 m depth.

There are scattered submerged flat boulders and

several emergent rounded ones that give the locality

its name and where waves break continuously.

A band of red turf algae, bleached during summer,

consisting predominantly of Chondrophycus

tumidus, Polyopes constrictus, and Gelidium australe

occurred in the sublittoral fringe zone as defined by

Womersley (1981), with sporadic denuded plants of

the fucoid Cystophora intermedia.

The sublittoral canopy layer with up to 90% cover

was dominated by patches of the fucoid Acrocarpia

paniculata and the spinous form of the laminarian,

Ecklonia radiata. Prominent but sporadic patches of

green Caulerpa flexilis var. muelleri and C. brownii

also occurred. Wet weights of algae collected during

2004 at 5 and 8 m depths (Fig. 1) indicate increased

dominance of Ecklonia and decreased dominance of

Acrocarpia and Cystophora moniliformis with depth.

An understorey of encrusting coralline red algae

(not collected), other red algae and Caulerpa spp was

present.

2. “The Hump” (HU), adjacent to a SE headland of

high wave energy

This site has near-vertical, jointed granitic rock

walls to a depth of 5 m then a stepped slope of about

20° to a coarse sandy bottom at 25 m depth. The

region is subject to high wave energy, with

considerable surge even on relatively calm days. A

distinct band of red algal turf, bleached in upper parts

during summer, with some Ulva occurred in an

extended sublittoral fringe, the result of high wave

energy. Asparagopsis armata and mats of dark red

Gelidium australe were present with Pterocladia

lucida prominent in shallow water.

At about 5 m depth the articulated corallines

Cheilosporum sagittatum and Haliptilon roseum

occurred in patches and as an understorey to a dense

canopy of Acrocarpia paniculata and Ecklonia.

Larger clumps of the red algae Melanthalia concinna

and Ballia callitricha with the brown algae

Halopteris pseudospicata, Homoeostrichus sinclairii

and Lobospira bicuspidata also occurred.

2500 Se ]

= 2000 [__] 5m depth |

2 Hi sm depth

~ 1500

St |

g 1000 |

oD |

= 500 |

o- — ee =

Se 89 £2 § § gf 8.

2 6 of eo & © = = © x =

o> £o on i ae} SO := aD @

Oo 5 Qa a. 2 oy = nS ==

(oa oma ox See + AS 9° (7p) fob)

= 3 O + Qn © Qs os

Oo ® o nO & ome) 0} Qe

qa D> Ee > O Cc oe axe pom

O O€£ fe) Om o Qs.

x & > ©

O 6 o =

Lu iu Be oO

Fig. 1. Dominant algal species collected at “The Boulders” in 2004: differences with depth.

118

R. N. BALDOCK & H. B. S. WOMERSLEY

7000

5m depth

_~ 6000 15

: | m

= 5000

= 4000 25m

= 3000

= 2000

1000

0 _

& 8 E Zg

oo 2 0 So

| as xd

Oo 0 o oO x

oc ee | oO ©

aS O= LU

Oo & oc

qt a 20

o oO

Fig. 2. Canopy species collected at “The Hump” in 2004: changes with depth.

250

E 200 5m

—

= 150 mm 15m

[o))

D |

= 50

0 | | — | —_ ——

£ 3 o 2 ae mets &

no © = = © a 2 a) cn

= 2 Onc a0 o> = ec Oo =

Go ec te’ 8 ES £ ©

5 2 7A} On 8-0 OH >

fe) = Qs 2". on noe)

12) o Ss) co) 2 oO fe)

xr ® oH oe Q a w

oO € £ Oo

a. je) ac

Fig. 3. Understorey algae collected at “The Hump” in 2004:

The brown canopy species and articulated

corallines persisted to 25 m deep, where the deeper-

water brown algae Scytothalia dorycarpa,

Glossophora nigricans, Carpoglossum confluens and

Cystophora platylobium occurred sporadically. Wet-

weights of canopy species (Fig. 2) indicate a reverse

changes with depth.

relationship with depth to that found at “The

Boulders”, and may be related to the effect of the

higher wave energy and steeper slope at the site.

The red algae Phacelocarpus peperocarpos and

Plocamium preissianum became increasingly

common with depth. (See Fig..3.)

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 119

Fig. 4. “The Monument”. Photo: A. Hirst.

Fig. 5. Inshore of “The Monument” with bleached algal

turf in the lower intertidal zone.

3. “The Monument’(MO), a low rocky promontory

on the NW side of the Mooring Bay

This is a relatively flat promontory of jointed

granitic rock extending about 300 m from the base of

a calcarenite cliff (see Fig.4). It is almost bisected

halfway along its length, and there is a shallow

channel to 3 m deep on its eastern outer end. There is

moderate wave action near the cliff face, with some

sand scour near the shore.

The shallow channel is subject to high wave surge

and the seaward extremity of the promontory

experiences high wave energy.

The different conditions along the length of the

promontory are reflected in the change of algal

dominants.

Inshore, calcified Liagora harveyana occurred in

shallow calmer water in gutters formed in the jointed

rock, with Cladosiphon vermicularis in the lower

eulittoral on shaded walls or in rock pools.

Cartilaginous red algae Chondrophycus tumidus and

Gigartina densa formed a distinct band, bleached in

summer, in the sublittoral fringe zone with some

articulated coralline algae (Fig. 5).

Cystophora moniliformis was prominent in shallow

seaward areas subject to wave surge, with Caulerpa

brownii in the lower eulittoral — upper sublittoral and

some Caulerpa papillosa.

120 R. N. BALDOCK & H. B. S. WOMERSLEY

Ecklonia radiata occurred in the sublittoral, but

had shorter stipes and was less dominant than at

other sites. Large patches of co-dominant Lobospira

bicuspidata and Cystophora monilifera occurred

mixed with some Acrocarpia paniculata. Patches

about 2 m in diameter of Caulerpa flexilis var.

muelleri, sometimes mixed with Caulerpa brownii

and Caulerpa geminata, occurred on horizontal

rocks 1 —4 m deep midway along the promontory.

Caulerpa obscura appeared in crevices and shaded

slopes almost to the low water mark, and

Dictyopteris muelleri was found at the entrance of

caves, 2 — 3 m deep.

The farthest extent of the promontory, exposed to

higher wave energy, had a prominent sublittoral

fringe with Ulva australis, Gelidium australe, and

denuded Cystophora intermedia.

4. Mooring Bay (MB)

This is a concave, sandy bay about 300 m across

bounded in the east by “The Boulders” and in the

north by “The Monument”.

A flat, granitic substratum partly sand covered with

a mono-specific Liagora harveyana community

occurred at 1 — 2 m depth on the eastern side.

Shepherd & Brook (2005) reported it to be sheltered,

with smooth granite boulders of 1 —2 m relief, depth

1 — 3 m with Cystophora and Sargassum spp and

Liagora, and patches of the articulated corallines

Corallina officinalis and Jania verrucosa.

5. Sea cave (SC), SE coast

A fracture in the granitic basement rock has been

eroded to a deep channel in the cliff face some 50 m

long and 15 m deep underwater, narrowing to a cave

11 m deep underwater ending abruptly at a rock fall

7 m deep underwater with a hole in the slumped

calcarenite ceiling above water (see Fig. 6). Near-

vertical sides and cave environment supported deep-

water and shade-loving bryozoans, hydroids and

algal species. These latter included the red

algae Rhodophyllis membranacea, Carpopeltis

phyllophora, Halymenia plana and Sonderopelta

coriacea, and the brown alga Homoeostrichus

sinclairit.

Western Isles (WI)

A string of five isles about 10 m high on the

exposed western side of the main island have been

formed by erosion of an amphibolite dyke (Robinson

Fig. 6. Sea cave entrance, with the jointed basement granitic rock overlain by slumped calcarenite.

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

et al. 1996). The calcarenite topping has been eroded

extensively leaving mainly conical granitic mounds

above water. Shepherd (pers. comm.) who also dived

at the site reports the sea floor to be generally 15 —

25 m deep, with moderate to strong surge, smooth

granite substratum, | — 2 m relief and in some places

undercut with caves. He noted that Ecklonia,

Acrocarpia and Cystophora spp formed the canopy

TABLE |. Site Codes used for algal collections.

121

layer. A form of Cystophora retorta with narrow

branches was collected.

NW coast of the main island.

This region is about 100 m offshore from “The

Monument”.

Collections at 10, 16, and 23 m were made by

divers on 6 February, 2004.

Code Site Depth (m) Collector(s) recorded Date

on herbarium sheets

Collections prior to 2004

NS __ N side of Althorpe I., SAORI expedition 9-13 R. Baldock 04.1.1964

EX coast exposed to high wave energy 5, 10, 15 J. Baker and K. Edyvane 26.x.1993

SH sheltered coast 5, 10, 15 J. Baker and K. Edyvane 27.x.1993

IT intertidal/drift collection A. Gaut 27-8.ix.2001

SS saline soak, S of “the Hump”, 50m inland E. Lawler 28ix.2001;

28.xi.2004

2004 expedition

BO “The Boulders” 0-8 R. Baldock 31.1.2004

HU = “The Hump” 4-25 R. Baldock 31.1.2004

MO “The Monument” 0-4 R. Baldock 01.11.2004

SC Sea cave 7-11 R. Baldock 03.11.2004

NW _ Northwest coast 10, 16, 23 J. Brook and R. Lewis 05.11.2004

Wi Western Isles 10, 27 J. Brook, R. Lewis and 06.11.2004

N. Barrett

MB_ Mooring Bay 1-2 S. Shepherd 11.11.2004

TABLE 2. List of Species with Depths in metres

Cyanophyta

Rivulariales-Rivulariaceae

Calothrix ?

Rivularia firma Womersley

Chlorophyta

Ulvales-Ulvaceae

Blidingia marginata (J. Agardh) P. Dangeard

tEnteromorpha clathrata (Roth) Greville ?

¥ Enteromorpha intestinalis (Linnaeus) Link?

T Ulva australis Areschoug

Cladophorales-Cladophoraceae

Apjohnia laetevirens Harvey

Chaetomorpha aerea (Dillwyn) Kitzing

Cladophora valonioides Sonder

Codiales-Codiaceae

Codium pomoides J. Agardh

Caulerpales-Caulerpaceae

Caulerpa brownii (C. Agardh) Endlicher

Caulerpa flexilis Lamouroux

Caulerpa geminata Harvey

Caulerpa obscura Sonder

Caulerpa papillosa J. Agardh

Caulerpa scalpelliformis (R. Brown) C. Agardh

Caulerpa vesiculifera Harvey

IT, on Chaetomorpha aerea

MO, mid-eulittoral

IT, in shallow pool

BO, MO, 0-3m

BO, 8m

IT, in shallow pool

HU, 17-25m

MO, Im

BO, MO, NS, SH, 0-13m

MO on Ecklonia base, 0-3m

IT, as drift; MO, NS, SH, 0-13m

BO, MO, SH, 0-10m

NS, NW, MO, 0-23m

NS, 9-13m

T the recent merging of Enteromorpha with Ulva by Hayden, H. S. et al. (2003) has not been recognized

122 R. N. BALDOCK & H. B. S. WOMERSLEY

TABLE 2. Cont.

Phaeophyta

Chordariales-Ralfsiaceae

Ralfsia verrucosa (Areschoug) J. Agardh

Chordariaceae

Cladosiphon vermicularis (J. Agardh) Kylin

Sphacelariales-Sphacelariaceae

Sphacelaria novae-caledoniae Sauvageau

Stypocaulaceae

Halopteris paniculata (Suhr) Prud’homme van Reine ?

Halopteris pseudospicata Sauvageau

Phloiocaulon spectabile Reinke

Cladostephaceae

Cladostephus spongiosus (Hudson) C. Agardh

Dictyotales-Dictyotaceae-Dictyoteae

Dictyopteris muelleri (Sonder) Reinbold

Dictyota diemensis Kiitzing

Dilophus fastigiatus (Sonder) J. Agardh

Glossophora nigricans (J. Agardh) Womersley

Pachydictyon paniculatum (J. Agardh) J. Agardh

Zonarieae

Distromium flabellatum Womersley

Distromium multifidum Womersley

Exallosorus olsenii (Womersley) Phillips

Homoeostrichus sinclairii (Hooker & Harvey) J. Agardh

Lobospira bicuspidata Areschoug

Zonaria spiralis (J. Agardh) Papenfuss

Zonaria turneriana J. Agardh

Sporochnales-Sporochnaceae

Sporochnus radiciformis (R. Brown) C. Agardh

Scytosiphonales-Scytosiphonaceae

Colpomenia sinuosa (Mertens) Derbes et Solier

Scytosiphon lomentaria (Lyngbye) Link

Laminariales-Alariaceae

Ecklonia radiata (C. Agardh) J. Agardh

Fucales-Hormosiraceae

Hormosira banksii (Turner) Decaisne

Seirococcaceae

Scytothalia dorycarpa (Turner) Greville

Cystoseiraceae

Acrocarpia paniculata (Turner) Areschoug

Carpoglossum confluens (R. Brown) Kiitzing

Caulocystis cephalornithos (Labillardiere) Areschoug

Cystophora intermedia J. Agardh

Cystophora monilifera J. Agardh

Cystophora moniliformis (Esper) Womersley & Nizamuddin

Cystophora platylobium (Mertens) J. Agardh

Cystophora retorta (Mertens) J. Agardh ?

Cystophora siliquosa J. Agardh

Sargassaceae

Sargassum paradoxum (R. Brown ex Turner) Hooker & Harvey

Sargassum vestitum (R. Brown) C. Agardh

Sargassum: Arthrophycus? (base only)

MO, on Cellana shell, lower eulittoral

MO, 0-3m

BO, on Metagoniolithon radiatum, 8m

HU, 0-4m

BO, MO, NW, 0-16m

HU, 17-25m

MO, 0-3m

MO, 1.5m

MO, NS, 0-13m

NS, NW, WI, 9-16m

HU, SC, WI, 7-27m

BO, IT, MB, NS, SH, 0-13m

IT as drift, SH, 10m

SH, 0-10m

SC, 7-1 1m

BO, EX, HU, NS, NW, SH, WL 5-27m

HU, MO, NS, 0-13m

BO, SH, 8-25m

HU, WI, 17-27m

NS, 9-13m

BO, MO, 1-8m

MO, and widespread but not collected

MO, 0-3m

HU, 17-25m

BO, MO, NS, 1-13m

HU, NS, WI, 9-27m

IT, as drift

BO, IT as drift, MO, 0-1m

BO, MO, 1-3m

HU, NS, 17-25m

WI, 10m (form with narrow branches)

MO, 3m

IT, as drift

BO, 0-5m

BO, MO, 1-8m

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

TABLE 2. Cont.

123

Rhodophyta

Balliales’-Balliaceae

Ballia callitricha (C. Agardh) Kiitzing

Nemaliales-Liagoraceae

EX, HU, IT as drift, NS, NW, WI, 5-27m

Helminthocladia australis Harvey

Liagora harveyana Zeh

Gelidiales-Gelidiaceae

Gelidium australe J. Agardh

Pterocladia lucida (Turner) J. Agardh

Pterocladiella capillacea (Gmelin)

Santelices & Hommersand

Gigartinales-Peyssonneliaceae

Peyssonnelia capensis Montagne

Peyssonnelia foliosa Womersley

Peyssonnelia novae-hollandiae Kiitzing

Sonderopelta coriacea Womersley & Sinkora

Polyidaceae

Rhodopeltis australis Harvey

Halymeniaceae

Carpopeltis phyllophora

(Hooker & Harvey) Schmitz

Halymenia plana Zanardini

Kallymeniaceae

Austrophyllis alcicornis (J. Agardh)

Womersley & Norris?

Callophyllis lambertii (Turner) J. Agardh

Cirrulicarpus nanus (J.Agardh) Womersley

Gigartinaceae

Gigartina brachiata Harvey?

Gigartina densa Edyvane & Womersley

Polyopes constrictus (Turner) J. Agardh

Dicranemataceae

Peltasta australis J. Agardh

Areschougiaceae

Callophycus laxus (Sonder) Silva

Callophycus oppositifolius (C. Agardh) Silva

Plocamiaceae

Plocamium angustum (J. Agardh)

Hooker & Harvey

Plocamium cartilagineum (Linnaeus) Dixon

Plocamium costatum (C. Agardh)

Hooker & Harvey

Plocamium dilatatum J. Agardh

Plocamium leptophyllum Kiitzing ?

Plocamium preissianum Sonder

Phacelocarpaceae

Phacelocarpus apodus J. Agardh

Phacelocarpus peperocarpos (Poiret)

Wynne, Ardre & Silva

Nizymeniaceae

Nizymenia conferta (Harvey)

Chiovitti, Saunders & Kraft

MB, MO, Im

MB, MO, 0-3m

BO, EX, MO, NS, NW, SH, WI, 0-16m

EX, HU, NW, SC, WI, 5-16m

IT in pool, SH, 0-5m

EX, WI, 15-27m

EX, 15m

EX, 10-15m

EX, HU, SC, WI, 7-27m

WI, 27m

EX, HU, NS, SC, WI, 5-27m

SC, 7-11MSC 7-11m

NW, 23m

NS, NW, WI, 9-16m

SH, ?m

MO, 0-3m

BO, EX, HU, MO, 0-10m

WI, 27m

WI, 10m

BO, EX, MO, NS, NW, SH, 0-23m

EX, IT, as drift; SH, 0-15m

EX, SH, 5-15m

WI, 27m

SH, 5m

EX, HU, NW, SH, 5-23m

HU, MO, NS, NW, 0-27m

EX, HU, NW, SC, SH, WI, 5-27m

NS, 9-13m

‘ tentatively placed in this systematic position on the basis of genetic sequencing by Choi et al. (2000)

124 R. N. BALDOCK & H. B. 8S. WOMERSLEY

TABLE 2. Cont.

Cystocloniaceae

Rhodophyllis membranacea (Harvey)

Hooker & Harvey ex Harvey

Rhodophyllis multipartita Harvey

Hypneaceae

Hypnea ramentacea (C. Agardh) J. Agardh

Hypnea valentiae (Turner) Montagne

Mychodeaceae

Mychodea marginifera (Areschoug) Kraft

Gracilariales-Gracilariaceae

Curdiea angustata (Sonder) Millar

Curdiea obesa (Harvey) Kylin

Melanthalia abscissa (Turner)

Hooker & Harvey

Rhodymeniales-Champiaceae

Champia zostericola (Harvey)

Reedman & Womersley?

Rhodymeniaceae

*Rhodymenia foliifera Harvey

Rhodymenia verrucosa Womersley

Corallinales-Sporolithaceae

Sporolithon durum (Foslie)

Townsend & Woelkerling.

Corallinaceae

Arthrocardia flabellata (Kitzing) Manza

ssp. australica Womersley & Johansen

Cheilosporum sagittatum (Lamouroux)

Areschoug

Corallina officinalis Linnaeus

Haliptilon roseum (Lamarck)

Garbary & Johansen

Jania affinis Harvey

Jania micrarthrodia Lamouroux

Jania verrucosa Lamouroux

Metagoniolithon radiatum (Lamarck) Ducker

Metagoniolithon stelliferum (Lamarck)

Weber-van Bosse

Metamastophora flabellata (Sonder) Setchell

Synarthrophyton patena

(Hooker & Harvey) Townsend

Bonnemaisoniales-Bonnemaisoniaceae

Asparagopsis armata Harvey

Delisea hypneoides Harvey

Delisea pulchra (Greville) Montagne

Leptophyllis conferta

(R. Brown ex Turner) J. Agardh

Ptilonia australasica Harvey

Ceramiales-Ceramiaceae-Wrangelicae

Wrangelia nobilis Hooker & Harvey

Wrangelia plumosa Harvey

Dasyphileae

Muellerena wattsii (Harvey) Schmitz

SC, 7-11m

HU, NW, WI, 10-27m

BO, NS, NW, 0-16m

MO, 0-3m

SC, 7-1 1m

WI, 27m

EX, MO, NS, HU, 0-15m

BO on Laurencia in lower eulittoral,

EX, HU, NW, SH, 5-25m

EX, MO, 15-16m

BO, 4-5m

MO, 0-3m

BO, EX, HU, NS, NW, SC, 4-16m

BO, MB, 0-1m

BO, on Ecklonia bases and Acrocarpia,

EX, HU, MO, NW, SH, 0-25m

WI, on Ballia, 27m

MO, 0-3m

MB, SH, 1-10m

BO, EX, MO, NS, 0-15m

SH, 10m

HU, SC, WI, 7-27m

WI, on Ballia and Phacelocarpus, 27m

BO, MO, SH, 0-S5m

HU, NS on Acrocarpia paniculata, 5-27m

NW, WI, 23-27m

WI on Ballia, 27m

WI, 27m

NW, 16m

IT, eulittoral pool

HU on Phacelocarpus peperocarpos, 15m

* The earliest name available for this complex, previously known as R. australis Sonder or R. sonderi Silva. (H.B.S.Womersley)

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

TABLE 2. Cont.

125

Antithamnieae

Acrothamnion preissii (Sonder) Wollaston

Antithamnion biarmatum Athanasiadis

Antithamnion hanovioides (Sonder) De Toni

Pterothamnieae

Inkyuleea mariana Choi, Kraft & Saunders

Heterothamnieae

Elisiella arbuscula (J. Agardh) Womersley ?

Griffithsieae

Griffithsia elegans Baldock

Ptiloteae

Euptilota articulata (J. Agardh) Schmitz

Spyrideae

Spyridia dasyoides Sonder

Ceramieae

Centroceros clavulatum (C. Agardh) Montagne

Ceramium filiculum Womersley

Ceramium pusillum Harvey

Dasyaceae

Dasya baldockii Parsons & Womersley

Heterosiphonia gunniana (Harvey) Reinbold

Heterosiphonia microcladioides

(J. Agardh) Falkenberg

Thuretia quercifolia Decaisne

Delesseriaceae-Nitophylloideae

Acrosorium ciliolatum (Harvey) Kylin

Crassilingua marginifera (J. Agardh) Papenfuss

Haraldiophyllum erosum (Harvey)

Millar & Huisman

Hymenena multipartita

(Hooker & Harvey) Kylin

Rhodomelaceae-Polysiphoniae

Polysiphonia decipiens Montagne

Polysiphonia sertularioides

(Grateloup) J. Agardh

Herposiphonieae

Herposiphoniella plurisegmenta Womersley

Lophothalieae

Haplodasya urceolata (Harvey) Parsons

Amansieae

Osmundaria prolifera Lamouroux

Laurencieae

Chondrophycus tumidus (Saito & Womersley)

Garbary & Harper

Laurencia clavata Sonder

Laurencia elata (C. Agardh) Hooker & Harvey

Laurencia filiformis (C. Agardh) Montagne

Laurencia majuscula (Harvey) Lucas

Laurencia spp

HU on Zonaria, W1 on Inkyuleea, 17-27m

SC on Metamastophora, 7-\\m

BO, MO on Polyopes constrictus, 0-3m

SC, WI, 7-27m

BO on Acrocarpia, 0-1m

SC on Glossophora, 7-\\m

NW, SC, WI, 7-23m

WI, 10-27m

IT rockpool

BO on Acrocarpia, 0-1m

BO on Acrocarpia, MO on Lobospira, 0-3m

NS, 9-13m

HU epiphytic, 17-25m

NW, 23m

HU on Homoeostrichus sinclairii,

SC on Glossophora bases , 5-1 1m

NW on Callophyllis, 16-23m

WI on /nkyuleea, 27m

NW on Haliptilon, 16m

BO on Acrocarpia, MO, 0-3m

MO, 0-3m

NS on Prerocladiella capillacea, 9-13m

BO on Cystophora intermedia, 0-1m

NS, 9-13m

BO, MO, 0-3m

BO, 5m

EX, 5m

IT as drift, SH, 5m

SH, 10m

IT, MO, 0-3m

126 R.N. BALDOCK & H. B. S. WOMERSLEY

Species for which the Island is the type locality:

Dasya_ baldockii, and ~~ _Herposiphoniella

plurisegmenta.

Extension of Distribution Range

The distributions for Peltasta australis and

Antithamnion biarmatum, previously known mainly

from Tasmania and Victoria, have been extended

westwards.

Discussion

The Althorpes are characterised by crystalline

granitic substrates, temperate waters and high wave

energy.

Canopy species of subtidal communities are

largely Acrocarpia paniculata and Ecklonia radiata,

with Cystophora monilifera and C. moniliformis also

prominent. Large patches of Caulerpa spp. and

Phacelocarpus spp. predominate at depth and as the

understorey.

Shepherd & Sprigg (1976) and Edyvane & Baker

(1998), believe the high macroalgal diversity and

productivity of islands at the entrance to South

Australian gulfs including the Althorpes, are

probably a result of a number of geographic factors

including high wave energy, summer nutrient

upwelling and geographic position between oceanic

and gulf waters with appreciable tidal flows from the

gulf. Edyvane & Baker pointed to the unique

position of the Gambier and Althorpe islands at the

boundary of autumn and summer water masses at

gulf entrances (Petrusevics 1993). This higher algal

diversity is, however, typical of the whole west coast

of Eyre Peninsula and south coast of Yorke

Peninsula.

Phillips (2001) added habitat hetero-geneity,

marine transgressions and regressions and lack of

mass extinctions to the list of factors contributing to

the high macroalgal biodiversity and endemism of

southern Australian regions.

Womersley & Edmonds (1958) had earlier

classified the South Australian coast on the basis of

intertidal and upper subtidal organisms (mainly

algae), wave action and substrate. Two categories of

coasts are relevant. One with extreme wave action

and steeply sloping crystalline rocks occurs on the

southwest tip of Yorke Peninsula, and west and south

Kangaroo Island. The other lies between areas of the

first and has more sheltered conditions, moderate

wave action, and flat calcarenite shore platforms

interspersed with sandy beaches, It is prominent on

southern Yorke Peninsula and the north coast of

Kangaroo Island. The Althorpes appear to lie

geographically at the junction of these two types, but

ecologically with the first.

The Althorpe Islands are positioned at the

boundary of the St Vincent Gulf and Eyre Bioregions

(as classified by IMCRA Technical Group 1998),

and contain features of both bioregions according to

Baker (2005). Edyvane & Baker (1998) noted the

higher species richness of the region’s outlying

islands, particularly Haystack Island. However, this

high diversity is typical of the high wave energy

coasts of Yorke and Eyre Peninsulas and Kangaroo

Island in general (see Womersley & Edmonds 1958).

Although only 144 species are documented in this

present paper, further detailed collecting in the

intertidal and sublittoral throughout the year would

undoubtedly increase the known flora from this

algal-rich southern Yorke Peninsula region.

Acknowledgements

SARDI financed the expedition, and provided the

research vessel Ngerin and diving facilities. A. Hirst

helped crop algae for biomass determinations and

provided photographs.

The Friends of Althorpe Is kindly hosted research

teams, and they and the Innes National Park staff

helped greatly with the collection and transfer to

shore of equipment and supplies. The State

Herbarium provided research facilities including

curatorial support. In particular, Ms Carolyn Ricci

was invaluable in collating and documenting

collections.

References

Baker, J. (2005). “Towards a System of Ecologically

Representative Marine Protected Areas in South

Australian Marine Bioregions — Technical Report”.

Report for Coast and Marine Conservation Branch,

Department for Environment and Heritage, South

Australia. 1250p.

Cnot, H-G., Krart, G. T. & SAUNDERS, G. W. (2000).

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(Rhodophyta): proposal of the Balliales ord. nov.,

Balliales fam. noy., Ballia nana sp. nov. and Inkyuleea

gen. nov. (Ceramiales). Phycologia 39:272-287

EpyvANE, K. S. & Baker, J. L. (1996). “Marine

Biogeography of Spencer Gulf, South Australia”. Final

report to the Nature Conservation Agency. (S. Australian

Research and Development Institute (SARDI) Aquatic

Sciences, West Beach, S. Australia.) 29pp +

& (1998) “Marine benthic survey

of Investigator Strait-Gambier Isles, South Australia”.

Report to Environment Australia (Marine Protected

Areas Program) Project D801, stage 4. 23pp +

(SARDI, Aquatic Sciences, West Beach, South

Australia.)

MARINE BENTHIC ALGAE OF THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 127

Haypen, H. S., BLomster, J., Macas, C. A., Siva, P. C.,

STANHOPE, M. J. & WAALAND, J. R. (2003). Linnaeus was

right all along: Ulva and Enteromorpha are not distinct

genera. Eur. J. Phycol. 38: 277-294.

IMCRA TecHNICAL Group (1998). Interim Marine and

Coastal Regionalisation for Australia: an ecosystem-

based classification for marine and _ coastal

environments, Version 3.3. Environment Australia,

Canberra.

Petrusevics, P. M. (1993). SST fronts in inverse estuaries,

South Australia: Marine Biogeography. Consultancy

Report to PISA, S. Australia. (SARDI, West Beach, S.

Australia.)

Puitiips, J. A. (2001). Marine macroalgal biodiversity

hotspots: why is there high species richness and

endemism in southern Australian marine benthic flora?

Biodiv. and Cons. 10:1555-1577.

RosINson, T., CANTY, P., Mooney, T. & Ruppuck, P.

(1996). “South Australia’s offshore islands”. Dept. of

Environment and Natural Resources South Australia.

SHEPHERD, S. A. & BROOK, J. B. (2005) Foraging ecology of

the western blue groper, Achoerodus gouldii, at Althorpe

Islands, South Australia. Trans. R. Soc. S. Aust. 129,

202-208.

SHEPHERD, S. A. & SpricG, R. C. (1976). Substrates,

sediments and subtidal ecology of Gulf St Vincent and

Investigator Strait. Jn Twidale, C & Webb, B. P. (Eds.)

“Natural History of the Adelaide Region.” pp. 161-174.

(R. Soc. S. Australia, Adelaide.)

WENIU, C., SCHAHINGER, R. B. & LENNON, G. W. (1990).

Layered models of coastal upwelling: a case study of the

South Australian region. Jn Davies, A. M. (Ed.)

“Modelling Marine Systems” pp. 73-91. (CRC Press,

Boca Raton, Florida.)

Womers_Ley, H. B. S. (1981) The marine ecology of

temperate coasts. Jn Clayton M.N. & King, R. J. (Eds),

“Marine Botany: an Australasian Perspective”, Ch. 11,

pp. 294-306. (Longman Cheshire, Melbourne.)

(1984) “The Marine Benthic Flora of Southern

Australia. Part I” (Govt. Printer, Adelaide).

(1987) “The Marine Benthic Flora of Southern

Australia. Part II” (Govt. Printer, Adelaide).

(1994) “The Marine Benthic Flora of Southern

Australia. Rhodophyta - Part IIA” (ABRS, Canberra).

(1996) “The Marine Benthic Flora of Southern

Australia. Rhodophyta - Part IIIB” (ABRS, Canberra).

(1998) “The Marine Benthic Flora of Southern

Australia. Rhodophyta - Part IIIC” (State Herbarium of

South Australia).

(2003) “The Marine Benthic Flora of Southern

Australia. Rhodophyta - Part IID” (ABRS, CANBERRA).

& Epmonps, S. J. (1958). A general account of

the intertidal ecology of South Australian coasts. Aust. J.

Mar. Freshw. Res. 9 (2):217-260, plates 1-12.

Transactions of the Royal Society of S. Aust. (2005), 129(2), 128-144.

SUBTIDAL MACROFLORA OF

ALTHORPE AND HAYSTACK ISLANDS, SOUTH AUSTRALIA

by J. L. BAKER!, G. J. EpGAR? & N. S. BARRETT?

Summary

Baker, J. L., EpGar, G, J. & BARRETT, N. S. (2005), Subtidal macroflora of Althorpe and Haystack Islands,

South Australia. Trans. R. Soc. S. Aust. 129(2), 128-144, 30 November, 2005.

The benthic macroflora at sites around the Althorpe and Haystack Is was surveyed in October 1993 and

January 2004. Nearshore reefs were dominated by the kelp Ecklonia radiata, the fucoid Acrocarpia paniculata

and mixed Cystophora species, whereas reefs at Haystack I. were dominated by a multi-species canopy of mixed

Sargassum and Cystophora species, with Ecklonia and Seirococcus axillaris also present. On sandy bottoms off

Althorpe I. and Haystack I., seagrass communities comprised mixed beds of up to five seagrass species, within

four genera (Amphibolis, Halophila, Posidonia, and Heterozostera). Biomass and species composition of the

canopy macroalgae were similar at those sites, which were surveyed in both 1993 and 2004, particularly at sites

where few species dominated the canopy. For the upper mid-sublittoral samples at two sites around Althorpe L.,

a similar average number of understorey species was recorded per m? in both 1993 and 2004. At comparable

sites, the understorey species composition differed between the two time periods; however, Caulerpa flexilis was

a dominant component during both 1993 and 2004. The islands are species-rich, with 267 species within

151 genera so far recorded to 27 m depth. During the 1993 survey, at least 180 species of macroalgae were

recorded in 29 quadrat samples, many of these from Haystack I. The macroal gal species richness at Althorpe and

Haystack Is is comparable with that of the Investigator Group and Nuyts Archipelago. These figures highlight

the importance of these islands in terms of macroalgal species richness in SA.

Key Worpbs: Macroalgae, subtidal survey, Althorpe Is, southern Australia, temperate macroalgal distributions,

macroalgal species richness.

Introduction

Macroalgae are a dominant feature of sublittoral

reefs in the temperate waters of southern Australia,

which is a particularly species-rich region

(Womersley 1984, 1987, 1990, 1994, 1996, 1998,

2003; Huisman ef al. 1998). In South Australia (SA),

which is especially rich in macroalgae, surveys at

several islands and bays during the past four decades

have contributed significantly to the knowledge of

macroalgal structure, distribution and zonation, and

the physical factors influencing those patterns (e.g.

Shepherd & Womersley 1970, 1971, 1976). This

paper adds to the catalogue of work from island

groups in SA coastal waters.

In 1993, reef macroflora and seagrass communities

at Althorpe and Haystack Is were sampled during a

state-wide benthic survey program (Edyvane &

Baker*). During the 2004 expedition some of the

South Australian Research & Development Institute, 2 Hamra

Ave, West Beach, South Australia 5024 and Dept Applied and

Molecular Ecology, University of Adelaide, Waite campus,

Urrbrae, South Australia 5064.

> Present address: 8 Fairfield Ave, Somerton Park, South Australia

5044, Email: jjbaker@senet.com.au

Tasmanian Aquaculture and Fisheries Institute, University of

Tasmania, Private Bag 49, Hobart, Tasmania 7011.

Edyvane, K. S. & Baker, J. L. (1998). “Marine Benthic Survey of

Investigator Strait — Gambier Isles, South Australia”. Report to

Environment Australia (Marine Protected Areas Program): Project

D801 (Stage 4). (SARDI Aquatic Sciences, South Australia).

same sites at Althorpe and Haystack Is were revisited

and sampled for biomass (Baldock & Womersley

2005). Further sampling of percent cover was

undertaken by the second and third authors during

surveys to assess the area for habitat value as part of

potential Marine Protected Areas. This paper

presents the results of those surveys from the two

time periods, particularly the data on biomass and

species richness, and considers the biogeographic

relations of the island group

Regional oceanography and site description

Frontal systems form seasonally in southern

Spencer Gulf, and the Althorpe Is are at the boundary

of these temperature fronts, where warmer gulf

waters and cooler deeper waters off western

Kangaroo I. meet, causing strong benthic

temperature and salinity differentials (Bruce & Short

1992; Petrusevics 1993). Sea surface temperatures

around the Althorpe Is are ~15 — 17° C in winter and

19 — 21° C in summer, and summer upwellings off

western Kangaroo I. and associated nutrients may

penetrate to the entrance of Investigator Strait

(Petrusevics 1993; Kampf et al. 2004). The

prevailing swell diffracts around Althorpe 1., such

that differences in water movement between various

sides of the island are slight (Shepherd ef a/. 2005).

The Althorpe Is lie at the boundary of the Eyre

(EYR) Bioregion and the Gulf St Vincent (GSV)

Bioregion, two of eight “meso-scale” biogeographic

SUBTIDAL MACROFLORA 129

Yorke Peninsula <>!

Ati. a 1

d me Zz

Nell / se)

e > _ Investigator Strait ee

| | ' :

= O-PS

Ne Wi pal ¢

- (a Kangaroo Island

Fig. 1. Map of the Althorpe and Haystack Is showing location in western Investigator Strait, at the boundary of the Eyre

and Gulf St Vincent Bioregions. Site numbers (see Table 1) are shown in white.

regions recognised for SA (IMCRA Technical Group

1998) (Fig. 1).

The Althorpe Is subtidally are of smoothly sloping

granitic basement reefs, interspersed with gaping

crevasses and chasms, except for the occasional

fallen calcarenite block in the shallows, whereas

Haystack I. has a substratum of eroded calcarenite

overlying a granitic spine. The geology and

topography of the Althorpe and Haystack Is are

described in some detail by Rankin ef al. (1991),

Murray-Jones & Shepherd (2005) and Zang (2005).

Methods

Sampling and analysis

1993: Sampling was undertaken from 24 — 31 Oct.

1993 with | m? quadrats. Mean % cover of

macroalgae and seagrasses was estimated visually,

For two sites each at Althorpe I. and Haystack I. (Fig.

1), the contents of four replicate quadrats (taken at

places which were representative of the habitat) at 5,

10, and on occasion 15 m depths were harvested, and

subsequently preserved, weighed and identified

(Table 1).

2004: Algal biomass data were collected at five

sites by Team A (Table 1, and see Baldock &

Womersley 2005), while Team B, as part of a wider

survey assessing biodiversity, used 20 replicate 50-

point quadrats (1 m2) placed along a series of 50 m

transects at 5 m and 10 m depths to estimate percent

cover at six sites (Table 1).

Multivariate classification procedures such as non-

metric multi-dimensional scaling (MDS) (Kruskal &

Wish 1978; Field ef al. 1982: Clarke 1993) and

cluster analysis (Romesburg 1984, Clarke 1993)

were used to elucidate spatial patterns in the 1993

and 2004 macroalgal biomass data. All weights given

are wet weights in g or kg m2. The data, which range

over four orders of magnitude, were transformed

(logi)) to increase the number of species that

contribute to the computation of dissimilarity

between samples, and to prevent the few species of

highest biomass from dominating analyses. Canopy

and understorey data were ordinated separately by an

MDS procedure in SYSTAT y. 11, using the Bray-

Curtis dissimilarity measure for the species by sites

matrices, and normalised Euclidean distance as the

constant in the Minkowski metric, for computing

distances between points in the MDS configuration.

Kruskal’s F-Stress Formula 1 (Kruskal & Wish 1978)

was used as the goodness-of-fit statistic for the

regression of 2-dimensional distances versus

dissimilarity values. The resulting positions in the

MDS plot were validated using a cluster analysis

(Romesburg 1984) procedure in SYSTAT vy. 1 1, with

the Bray-Curtis dissimilarity measure as a distance

metric, and the UPGMA clustering algorithm. Other

methods used to display patterns of sampled species

composition and/or abundance include summary

tables and charts, and a species accumulation curve.

Results

Macrofloral Composition

Ecklonia radiata is generally the dominant canopy

species, with Acrocarpia paniculata as a co-

dominant, on hard substrates in the mid-sublittoral (5

~— 15 m depth) around Althorpe I. (Table 2, Fig. 2A,

B), except near the sand line near seagrass beds (e.g.

10 m depth, Boulders - see Table 2). At Site 2,

130 JL. BAKER, G. J. EDGAR & N. S. BARRETT

TABLE 1. Sites sampled, and collecting methods used during surveys of Althorpe and Haystack Is in 1993 and 2004. Site

numbers are shown in Fig. 1, B= biomass in g wet weight m2. PC = percentage cover m?.

Team, Site Number Sampling Method

Place location, and depth and variable measured

1993 Site 1. The Boulders, 5, 10 m 1.0 m? quadrats B, PC

Althorpe I. Site 2. Off Chain Islet, 5, 10, 15 m

A-2004 Site 1, The Boulders, 0 — 8 m 0.25 m?— 1.0 m?

Althorpe I. Site 2. Off Chain Islet, 4-27 m quadrats

Salmon Inlet / Sea Cave, 7— 11m B

Site 3. NW Bay, 10, 16, 23 m

Site 5. Western Isles, 10, 27 m

B-2004 Site 1. The Boulders, 5, 10 m 50 points per quadrat,

Althorpe I. Site 2. Off Chain Islet, 5, 10 m 20 quadrats per site. PC

Site 3. NW Bay, 10 m

Site 4, Swallowtail Bay, 5, 10 m

Site 5, Western Isles 5, 10 m

1993 Site 6,5, 10m 1.0 m? quadrats. B, PC

Haystack I. Site 7, 10m

B-2004 Site 6,5 m; Site 7, 5m 50 points per quadrat,

Haystack I. 20 quadrats per site. PC

Ecklonia comprised 65 — 75% cover and ~2 —8 kg m?

from 5 mand 15 m depth in 1993, and similarly high

cover and biomass in 2004. Elsewhere, at Sites 1, 3,

and 4, biomass and % cover of Ecklonia were also

high in 1993 and 2004 (Table 2, Fig. 2A,B).

In the upper mid-sublittoral (5 — 10 m), the cover

and biomass of Acrocarpia were high at Site 2, with

values of 1.5 — 3.6 kg m? recorded in some quadrats.

At the same depths sampled off Haystack I., this

species was only a minor component of the canopy

flora. A few species of Cystophora, mainly C.

moniliformis and C. monilifera, were also present in

the mid-sublittoral around Althorpe [., although the

latter species was not recorded in 1993 at Site 2

(Table 2).

At Althorpe 1, during both the 1993 and 2004

surveys (Table 1), the fucoid Seirococcus axillaris,

was largely absent and, when present, occurred only

as a minor component of the canopy (i.e. average

<100 g m? at Site 2 at 15 m). In contrast, at Site 7,

Haystack [., Seirococcus was a major canopy species

at 10 m depth (Table 2), with 1.0 — 4.2 kg m?

recorded in 1993. The 2004 survey also recorded S.

axillaris at Site 6, Haystack I., but in low densities.

The fucoid Scytothalia dorycarpa (closely related

to Seirococcus) was uncommon or absent at most

sampled sites around Althorpe and Haystack Is in

1993 and 2004 (Table 2, Fig. 2A,B); however it was

recorded at 5 m depth at Site 5 (Fig. 2B), and at 23

m at Site 3, with a biomass of 960 g m>°.

Five species of seagrass were recorded on sand at

10 m depth at Site 1, Althorpe L, and Site 6,

Haystack I. (Table 2). At Site 1, Posidonia sinuosa

was dominant but P angustifolia was not recorded,

whereas at Site 6, the opposite was found. At both

sites Halophila australis, Heterozostera nigricaulis

(formerly H. tasmanica), and Amphibolis antarctica

were also recorded.

At Site 5, upper mid-sublittoral canopy

composition was similar to that recorded around

Althorpe L., with Ecklonia as the dominant cover, and

lesser quantities of Acrocarpia and species of

Cystophora (Fig. 2A,B). Differences in the canopy at

Site 5 included the presence of Scytothalia

dorycarpa and differences in the composition of the

Cystophora species present (e.g. C. retorta, recorded

at 5 m).

In deeper water (23 m) at Site 3, Acrocarpia

(1690 g m2), Scytothalia dorycarpa (960 g m”),

Seirococcus axillaris (865 g m*) and Cystophora

platylobium (670 g m) occurred among the

dominant Ecklonia ( ~10 kg m); however, samples

at 27 m depth at the same site revealed Ecklonia

(biomass 0.7 — 2 kg m7”) as the only large brown alga,

among a predominantly rhodophyte community

(biomass 800 — 1700 g m*). Common species were:

Delisea pulchra, Ballia callitricha, Inkyulea

mariana, Rhodophyllis multipartita, Phacelocarpus

peperocarpos, Sonderopelta coriacea, and Spyridium

dasyoides, and understorey brown algae such as

Homoeostrichus sinclairti and Zonaria spiralis.

Other red macroalgae recorded in the deeper waters

at Site 3 included Carpopeltis phyllophora,

Callophycus — oppositifolius, Curdiea obesa,

Leptophyllis conferta, Peltasta australis, Plocamium

dilatatum, Ptilonia australasica, Rhodopeltis

australis, the arborescent coralline Metamastophora

flabellata, the coralline Jania pulchella (= J. affinis)

SUBTIDAL MACROFLORA 131

Ecklonia

Acrocarpia

Cystophora monilifera

C. moniliformis

C. retorta

C. retroflexa

Sargassum sonderi

Sargassum fallax

Sargassum sp.

Ecklonia

Acrocarpia

Scytothalia dorycarpa

Seirococcus axillaris

Cystophora monilifera

Scytothalia dorycarpa

NW Haystack

NE Haystack

Swallowtail Bay

The Hump / off Chain Islet

The Boulders

Swallowtail Bay

NW Bay

The Hump / off Chain Islet

Western Isles

The Boulders

Fig. 2. Mean percentage cover of canopy macroalgae recorded at (A) 5m depth and (B) 10 m depth at selected sites around

Althorpe I. and Haystack I. in 2004. Site numbers and locations are described in Table 1.

and the small calcareous epiphyte Synarthrophyton

patens.

In contrast to Althorpe Is, Acrocarpia was

uncommon at Haystack I. and Ecklonia was co-

dominant at 5 — 10 m depth with other taxa including

Sargassum and Cystophora species and Seirococcus

axillaris (Table 2; Fig. 2A). Haystack I. was rich in

Sargassum species, with 10 species recorded from

the 1993 and 2004 surveys combined. The major

component canopy species were variously S.

linearifolium, S. verruculosum, S. sonderi, S. fallax,

C. moniliformis, and C. monilifera, with some

differences in relative abundance between the two

sampling periods (Table 2; Figs 2A, 4). At 10 m

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Data for 1993 are biomass (g wwt / m7’), averaged for replicate quadrat samples, with SD bars. Data for 2004 are

percentage cover. Similarity in canopy species composition between 1993 and 2004 is indicated by overlapping bars (1.c.

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Fig. 4. Non-metric multi-dimensional scaling and cluster analysis of the log-transformed biomass data for (A) canopy

species, and (B) understorey species, recorded at Site 1 (The Boulders) and Site 2 (The Hump) at Althorpe I. in 1993 and

2004. Numbers after the site name indicate depth (m) and year (93 or 04).

SUBTIDAL MACROFLORA 137

depth, canopy species recorded at Haystack I. that

were not common at other sampled sites included

Carpoglossum confluens and Seirococcus axillaris.

At the Althorpe I. sites, common understorey

species in the mid-sublittoral included: species of

Caulerpa (particularly C. flexilis); various turfing

browns; and numerous erect and encrusting red algae

(Table 2). During the 1993 survey, Caulerpa flexilis

was especially abundant at 5 m at Site | (Table 2).

Reefs in the upper mid-sublittoral Sites 6 and 7,

Haystack I., were particularly rich in understorey

species, whose species composition differed

considerably from that recorded at Althorpe I. (Table

2, Appendix 1).

Comparison of 1993 and 2004 Collections

The 2004 survey has shown some consistency over

time in the composition of the main canopy species

over the 11-year period, particularly the dominance

of Ecklonia and Acrocarpia on mid-sublittoral reefs

around Althorpe L, and the co-dominance of

Ecklonia amongst a mixed canopy of Cystophora and

Sargassum species on reefs around Haystack I.

(Table 2, Fig. 2); however, one noticeable difference

between the two sampling periods was the absence of

Acrocarpia in samples taken at Site | (5 m) in 1993,

and its dominance there in 2004. Haystack I. (Site 6)

also showed striking differences over the 11-year

period in species composition and abundance of

some species at 5 m, notably Sargassum (Table 2,

Fig. 3). Sargassum sonderi was the major species

recorded in 2004, whereas S. linearifolium, S.

paradoxum, S. spinuligerum and S. verruculosum

were recorded most frequently in 1993.

A multivariate analysis of the canopy and

understorey data for the two survey periods indicated

little difference between samples at 5 and 10 m

depths at Site 2 (off The Hump). In both 1993 and

2004 these pairs of data had a low calculated Bray-

Curtis distance, and thus appeared close together in

the ordination space and the cluster tree (Fig. 4A).

Similarly, the 10 m samples at Site 2 in 1993 and

2004, and the 5 and 8 m samples at Site 1 (The

Boulders) in 2004 were relatively close (Fig. 4A),

whereas the 5 m sample from 1993 was relatively

TABLE 3, Numbers of understorey species per m? recorded

in 1993 and 2004, at 5 m depth at Sites 1 and 2.

Understorey Site 2 Site |

No. m? (1993, 2004) 13, 14 21, 16

Sum of all species recorded 22 31

in 1993 and 2004

No. species >4 g m? recorded I 4

in both 1993 and 2004

Total no. and % of species recorded 6 6

in both 1993 and 2004 (27%) (19%)

isolated, primarily because this was the only

Althorpe I. site sample at which Acrocarpia

paniculata was not recorded.

For the understorey data, the most similar pairs of

samples were from 5 and 8 m depths at Site |

(sampled in 2004), and from 5 and 10 m at Site 2

(also 2004). Generally, the multivariate analysis

showed that samples from comparable depths taken

at the two different time periods were not similar,

with few 1993 and 2004 samples grouping closely

together in the ordination space.

The average number of understorey species

recorded per m? at 5 m depth at Sites 1 and 2 was

remarkably similar over time (Table 3), although the

species composition differed substantially (Table 2).

Thus, <30% of understorey species were common to

both sites at 5 m depth in both years (Fig, 2, Table 3).

Notable understorey species recorded during both

1993 and 2004 included Ballia callitricha at Site 2,

and Gelidium australe, Plocamium angustum and

Metagoniolithon radiatum at Site 1. For all samples

taken between 5 and 15 m deep at Site 2, 40% of the

understorey species were recorded during both time

periods.

Spatial Patterns in Species Composition and

Richness

After averaging replicate samples for each depth,

the number of canopy species per m? was strongly

correlated with the number of understorey species

per m? at Althorpe and Haystack Is sites (r = 0.79).

Where a dense cover of Ecklonia (and sometimes

Acrocarpia) were dominant (e.g. at 5 — 15 m depth at

Site 2), few other canopy species were present, and

fewer understorey species, compared with sites (e.g.

Sites 6, 7) where Ecklonia and/or Acrocarpia were

less dominant (Table 2).

The sampled species composition in the mid-

sublittoral differed both within and between islands.

At the finest scale (replicate quadrats) there were few

species in common between samples. For example,

of the 27 species recorded at 5 m at Site | in 1993,

about half were found only in one of four replicate

quadrats, and only four species (the common

Cystophora moniliformis, C. monilifera, Caulerpa

flexilis, and Plocamium angustum) were found in all

quadrats. Similarly at the 10 m site, about half of the

species were recorded in only one quadrat, and only

four of the 45 species were found in all four replicate

quadrats. At the 5 m site at Site 2, only three

(Ecklonia radiata, Ballia callitricha and Peyssonnelia

novae-hollandiae) of the 23 species were recorded in

three replicate quadrats. At Althorpe I., the

understorey species composition differed sub-

stantially between Sites 1 and 2 (Table 2; Fig. 4B:

Appendix). For example, during the 1993 survey,

only 12% of macroalgal species recorded on reefs at

138 J. L. BAKER, G. J. EDGAR & N.S. BARRETT

10 m deep were common to both sites. Of the 50

understorey species that were collectively recorded

from 5 — 10 m at the two sites in 1993, only 8 (16%)

were found at both sites. In the 2004 sampling, 27%

of the macroalgal species recorded at 5 m were

common to the two sites. Differences in species

composition were also observed between sites of

similar exposure at two different islands. For

example, only 18% of the species from 5 m depth

samples at Sites 1 and 6 were common to both.

Haystack I. was particularly rich in macroalgal

genera and species, both in the canopy and the

understorey, compared with Althorpe I. Using the

averaged biomass data (Table 2) for replicate quadrats,

2 — 6 canopy species per m? were recorded for the

Althorpe I. sites in 1993, compared with 13 at the

Haystack I. sites. Similarly, for understorey species,

<30 species per m* were recorded at most Althorpe I.

sites in 1993, compared with 49 — 67 at Haystack I.

(Table 2, Appendix). Within genera, examples of the

species richness at Haystack I. include the chlorophyte

Codium (4 species), and the rhodophytes Mychodea (5

species) and Hymenocladia (3 species). The family

Rhodomelaceae was well represented at Haystack I.,

with at least 15 species recorded during the 1993

survey (Table 2, Appendix 1).

The Althorpe Is and Haystack I. together were rich

in macroalgae, with at least 232 species within 136

genera recorded from the depth range 3 — 27 m. In

addition to those cited above for Haystack I., other

examples of the species richness at the Althorpe Is

include: the canopy genera Sargassum (10 species)

and Cystophora (7 species); the understorey genera,

Caulerpa (10 species), Dictyota (5 species),

Phacelocarpus (4 species), Plocamium (7 species),

Laurencia (5 species), Griffithsia (3 species) and

Heterosiphonia (3 species) (Table 2, Appendix).

While neither survey was designed to determine

species richness, it is notable that during the 1993

survey alone, at least 180 species of macroalgae were

recorded in 29 quadrat samples, from Sites 1 and 2,

Althorpe I., and Sites 6 and 7, Haystack I. (Table 2;

Fig. 5; Appendix). The “steps” in the curve of

cumulative species in Fig. 5 result from sampling at

species-rich Haystack I., where most of the 11 x 1m?

quadrat samples yielded species not previously

recorded in other samples. The curve in Fig. 5, which

does not reach an asymptote, shows that further

sampling from Haystack I. would likely yield even

more species of macroalgae.

Discussion

Macroalgal Biogeography of Althorpe and Haystack

Due to its central geographic location in the

Flindersian Province, the Althorpe Is contain many

species from the warmer western part of the

Flindersian Province, and the cool water Maugean

Sub-Province, as well as species with a broad

Flindersian distribution, extending to SE NSW

(Womersley 1990). The islands also lie between the

Eyre and Gulf St Vincent Bioregions, and have

various geomorphological, oceanographic and

biological features that are characteristic of each of

these bioregions. These include: the high species

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Fig. 5. Species accumulation curve for 29 x 1 m? quadrat samples from Althorpe and Haystack Is Samples were taken at

5, 10 and 15 m, at Sites 1,2 Althorpe L., and Sites 6,7 Haystack I. Samples are plotted in randomised order, and include

all canopy and understorey species (except for encrusting corallines).

SUBTIDAL MACROFLORA 139

diversity and abundance of red macroalgae, and the

seasonal influence of nutrient-rich, cooler water

upwellings, characteristic features of the Eyre

Bioregion (IMCRA Technical Group 1998, Baker

2005).

Ecklonia, Acrocarpia, and widely distributed

species of Sargassum and Cystophora together form

the major structural components of the reef system.

Examples of species with a western Flindersian

distribution (Cowan 2000) include the green

Cladophora valonioides, and the rhodophytes

Cliftonaea pectinata, Dasya_ baldockii, and

Heterosiphonia_ callithamnium. Maugean Sub-

Province species include the canopy brown

Sargassum vestitum, the turfing brown Exallosorus

olsenii, and the reds Gigartina muelleriana, Peltasta

australis, Ptilonia australasica, Muellerena wattsii,

Melanthalia abscissa, Shepleya wattsii, Mychodea

hamata, and Gloiocladia fruticulosa.

The canopy species composition recorded at

Althorpe and Haystack Is shares some common

features with canopies of island reefs both to the west

(e.g. Investigator Group off western Eyre Peninsula)

and east (e.g. West Island, Encounter Bay).

Similarities include the dominance of Ecklonia,

Acrocarpia paniculata, and the presence of mixed

Cystophora species (Shepherd & Womersley 1970,

1971; Baker & Edyvane 2003).

At Haystack I., the mixed canopy of various

Sargassum and Cystophora species, co-dominant

with Ecklonia, is a feature of other less wave-

exposed reefs throughout the central SA coast, such

as the Sir Joseph Banks Group, Spencer Gulf (JLB

unpublished data). The widely distributed

Flindersian species Scytothalia dorycarpa, and the

more easterly (Maugean) species Seirococcus

axillaris, were not common at the Althorpe Is or

Haystack I., although the former species is dominant

elsewhere in central SA, and the latter at islands

further east (e.g. West I.).

The records of six species in this survey extend

their known geographic range to the west. These

species, mostly Maugean, are: Exallosorus olsenii

(sub nom. Homoeostrichus olsenii in Womersley

1987), Sargassum vestitum, Cystophora retroflexa,

Gigartina muelleriana, Peltasta australis, and

Ptilonia australasica.

Also of note from these surveys are records of

species with limited known spatial distributions in

SA. Examples include: (i) the uncommon coralline

Amphiroa gracilis, a sub-tropical species known

from WA, SA and Qld (Cowan 2000). In SA it is

found mainly in the Gulfs region and around Yorke

and Eyre Peninsulas (Womersley 1996; Plant

Biodiversity Centre and DEH 2004); and (ii)

Arthrocardia wardii, an articulated coralline whose

distribution extends to NSW and Tas. In SA, the

species has been found only in Investigator Strait and

at Kangaroo I. (Womersley 1996; Plant Biodiversity

Centre and DEH 2004).

Canopy Species Composition in Space and Time

A number of studies have shown that on reefs

where Ecklonia is dominant, the corresponding

understorey is low in biomass and/or species due

inter alia to effects of shade, scour, and occupation

of available space (Kennelly 1987, 1988; Schiel

1990; Kendrick et al. 1999; Fowler-Walker &

Connell 2002; Turner & Cheshire 2003; Kendrick er

al, 2004). The Althorpe I. reefs were mostly

dominated by canopies of Ecklonia and Acrocarpia,

with few other canopy species, and a consequent low

diversity and abundance of understorey species.

The higher species richness at Haystack I. than at

Althorpe Is may be due to a combination of factors.

The highly rugose, actively eroding, calcareous

substrate at Haystack I., compared with the smooth

granite substrate at Althorpe Is, creates more micro-

habitats and greater opportunity for algal propagules

to settle, particularly within the Rhodophyta. Also,

the mixed Sargassum-Cystophora canopy at

Haystack I. promotes a more diverse understorey

(e.g. Kendrick et al. 2004), possibly through the

provision of more space and light, and less scour,

compared with canopies of Ecklonia. A feature of

increased species richness is the presence of many

rare or uncommon species (Hubbell 2001), as we

found at Haystack I.

Despite the fact that the exact sites were not

revisited in 2004, the apparent consistency over time

of canopy composition at these sites at Althorpe I. (a

feature previously noted by Baker & Edyvane (2003)

at Nuyts Archipelago) suggests a measure of stability

in these assemblages. Although alternative states in

algal assemblages may occur (Turner & Cheshire

2003), the prevailing exposure at any given site may

ensure the long-term persistence of particular canopy

genera (e.g. Fig. 4, Turner & Cheshire 2003).

At Haystack I. the main differences observed over

the 11 years were in the species and cover of

Sargassum at Site 6. These could be variously due to

shifts from one alternative state to another (see

above), and to seasonal differences in time of

sampling between surveys. As shown by Edgar

(1983), some Sargassum species seasonally shed

reproductive branches and laterals.

Understorey Species Composition in Space and Time

At sites investigated in this study, generally <30%

of the understorey species were recorded in both

1993 and 2004, although a similar number of

understorey species per m? were recorded during

both time periods. The apparent persistence of

Caulerpa flexilis at 5 m at Site 1 at both time periods

140 J. L. BAKER, G. J. EDGAR & N. S. BARRETT

was presumably facilitated by its stolonic matte,

which can pre-emptively occupy space and exclude

competitors (Shepherd 1981). Changes in species

composition over time are likely for many reasons.

These include: fine-scale patchiness of species;

rarity of many species, especially when species

richness is high (Baker & Edyvane 2003);

opportunistic and ephemeral strategies of some

species (Shepherd 1981); and effects of shading,

storms, grazers, sediment and scour (Kennelly 1987,

1988; Connell 2005).

Species Richness

Intertidal and subtidal collecting at Althorpe and

Haystack Is to 30 m depth have so far yielded a total

of 276 species within 151 genera. This compares

with 240 species in the Investigator Group (Shepherd

& Womersley 1971; Baker & Edyvane 2003), ~230

species at St Francis Isles (Shepherd & Womersley

1976; Womersley & Baldock 2003), 364 species at

Waterloo Bay (Shepherd & Womersley 1981), and

132 species at West I. (Shepherd & Womersley

1970). Hence the Althorpe and Haystack Is have a

similarly rich flora to other islands of the eastern

Great Australian Bight, which is notable for its algal

diversity. Thus, of the estimated 961 macroalgal

species recorded in SA waters (Womersley 1984,

1987, 1994, 1996, 1998, 2003; Cowan 2000), at least

28% of them, excluding the crustose corallines, have

been found to date at these islands.

Acknowledgments

We thank the many organisations and individuals

who assisted in these surveys and their supporting

organizations: the former Australian Nature

Conservation Agency, and the Australian Heritage

Commission for funding the 1993 survey; SARDI

for provision of MRV N¢gerin, and diving facilities in

1993 and 2004; R. N. Baldock for collecting and

processing algal biomass data in the 2004 expedition,

and for algal identifications; A. Bloomfield and

J. Brook for assistance in the collection of percent

cover and species data; the Marine Protected Area

program of Department for Environment and

Heritage for funding the visit and work of the second

and third authors; the Friends of Althorpe Islands for

hosting research teams in 2004; Innes National Park

staff for logistic assistance; the State Herbarium

for research facilities and curatorial support;

especially Professor H. B. S. Womersley for algal

identifications; K. Edyvane, the survey team leader

in 1993; divers assisting with algal collections,

namely G. Andrews, R. Baldock, J. Brook, B. Davies,

A. Hirst, M. Kinloch, R. K. Lewis, I. McGrath,

P. Preece, S. A. Shepherd, and G. Westphalen; the

crew of the MRV N¢eerin for their tireless assistance;

G. Lorenzin, A. Doonan, V. Day, S. Deakin and

C. Ricci for laboratory assistance; and lastly the

editors and anonymous reviewers for helpful

criticism and improvements to the paper.

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Appendix

List of sites at which minor species (<5 g m7?) were recorded at Althorpe and Haystack Is. A full list of species is available

from the first author. Names of collectors are given by Baldock & Womersley (2005).

Code Site Depth (m) Date

NS N side of Althorpe I. 9-13 4.1.1964

NW Site 3. Althorpe L. 10, 16, 23 4-11.11,2004

SE Site 2, Althorpe I. 5, 10, 15 26.x.1993, 31.i-11.ii1.2004

NE Site 1, Althorpe L. 5, 10 27.x.1993, 31.i1-4.11.2004

H1 Site 6, Haystack I. 5, 10 29.x.1993

H2 Site 7, Haystack I, 10 29.x.1993

MO The Monuments, Althorpe |. 0 - 4 1.11.2004

sc Sea Cave, Althorpe I. 7-11 3.i1.2004

142 J. L. BAKER, G. J. EDGAR & N.S. BARRETT

SW Site 4, Althorpe I. 5, 10 4-11.ii.2004

W Site 5, Western Isles. 5, 10, 27 4-11.ii.2004

Hl Site 6, Haystack I. 5, 10 29.x.1993; 4-11.ii.2004

H2 Site 7, Haystack I. 5, 10 29.x.1993; 4-11 .ii.2004

Chlorophyta

Caulerpales - Caulerpaceae

Caulerpa brownii (C. Agardh) Endlicher NE, MO, NS, SE, SW, H1, H2, 0-13m

Caulerpa scalpelliformis (R. Brown) C. Agardh W, NS, NW, NE, MO, H1, H2, 0-23m

Phaeophyta

Chordariales - Chordariaceae

Polycerea nigrescens (Harvey ex Kiitzing) Kylin H1, 10m

Dictyotales - Dictyotaceae - Dictyoteae

Dicytota alternifida J. Agardh NE, 10m

Pachydictyon polycladum (Kiitzing) Womersley H1, 5m

Dictyotales - Zonarieae

Dictyopteris muelleri (Sonder) Reinbold — MO, SE, 1.5-10m

Distromium flabellatum Womersley NE, H2, 10m

Distromium multifidum Womersley NE, 5m

Lobophora variegata (Lamouroux) Womersley ex Oliveira H1, H2, 5-10m

Zonaria angustata (Kiitzing) Papenfuss H1, H2, 5-10m

Zonaria crenata J. Agardh H2, 10m

Fucales - Cystoseiraceae

Myriodesma integrifolium Harvey H1, 5m

Rhodophyta

Gelidiales-Gelidiaceae

Gelidium crinale (Turner) Gaillon H1, 5m

Gigartinales - Gigartinaceae

Gigartina muelleriana Setch. & N.L. Gardner H2, 10m

Gigartina pinnata J. Agardh SE, H1, 5m

Polyopes constrictus (Turner) J. Agardh SE, NE, MO, 0-10m

Gigartinales - Dumontiaceae

Dudresnaya australis J. Agardh ex Setch. H2, 10m

Gigartinales - Kallymeniaceae

Polycoelia laciniata J. Agardh SE, 10m

Glaphrymenia pustulosa J. Agardh H2, 10m

Gigartinales - Areschougiaceae

Callophycus oppositifolius (C. Agardh) Silva NE, W, 10m

Erythroclonium angustatum Sonder H1, 10m

Rhabdonia coccinea (Harvey) Hooker & Harvey H2, 10m

Rhabdonia verticillata Harvey H1, 5m

Solieria robusta (Greville) Kylin H1, H2, 5-10m

Gigartinales - Plocamiaceae

Plocamium costatum (C. Agardh) Hooker & Harvey SE, NE, 5-15m

Gigartinales - Phacelocarpaceae

Phacelocarpus sessilis Harvey ex J. Agardh NE, 10m

Gigartinales - Nizymeniaceae

Nizymenia conferta (Harvey) Chiovitti, Saunders & Kraft NS, H1, H2, 5-13m

Gigartinales - Cystocloniaceae

Rhodophyllis membranacea (Harvey) Hooker & Harvey ex Harvey H1, SC, 5-11m

Rhodophyllis multipartita Harvey | SE, NW, W, 10-27m

Gigartinales - Mychodeaceae

Mychodea hamata Harvey H1, 10m

Mychodea marginifera (Aresch.) Kraft H1, 10m

Mychodea pusilla (Harvey) J. Agardh H1, 10m

SUBTIDAL MACROFLORA 143

Gracilariales - Gracilariaceae

Gracilaria cliftonii Withell, A. Millar & Kraft H1, 10m

Rhodymeniales - Rhodymeniaceae

Botryocladia sonderi Silva H1, 5m

Erythrymenia minuta Kylin H2, SE, 10-15m

Gloiocladia australis (J.Agardh) Norris H1, 10m

Gloiocladia fruticulosa (Harvey) Norris H2, 10ra

Gloiosaccion brownii Harvey H1, 10m

Hymenocladia chondricola (Sonder) Lewis H1, 10m

Hymenocladia divaricata Harvey H1, 10m

Hymenocladia usnea (R. Brown ex Turner) J. Agardh H1, 10m

Rhodymeniales - Champiaceae

Champia affinis (Hooker & Harvey) J. Agardh H1, 10m

Champia viridis C. Agardh NE, H1, H3, 5-10m

Rhodymeniales - Lomentariaceae

Lomentaria sp. H2, 10m

Corallinales - Corallinaceae

Amphiroa anceps (Lamouroux) Decaisne SE, NE, NW, H1, 5-15m

Amphiroa gracilis Harvey NW, SE, SW, 5-10m

Arthrocardia wardii (Harvey) Areschoug H1, 10m

Jania pulchella (Harvey) Johansen & Womersley NE, H1, 5-10m; W, on Ballia, 27m

Jania verrucosa Lamouroux NE, 1-10m

Corallinales - Mastophoroideae

Metamastophora flabellata (Sonder) Setchell SE, SC, W, 7-27m

Ceramiales - Ceramiaceae

Perischelia glomulifera (J. Agardh) J. Agardh = H2, 10m

Ceramiales - Ceramiaceae - Wrangelieae

Anotrichium “sp.1” NE, H2, 5-10m

Euptilocladia spongiosa Wollaston H2, 10m

Hirsutithallia formosa (Harvey) Wollaston & Womersley H1, H2, 10m

Involucrana crassa (Hooker & Harvey) Gordon-Mills H2, 10m

Shepleya wattsii (Harvey) Gordon-Mills © H2, 10m

Wrangelia nobilis Hooker & Harvey NW, 16m

Wrangelia velutina (Sonder) Harvey H1, 10m

Ceramiales - Ceramiaceae - Crouanieae

Ptilocladia australis (Harvey) Wollaston H1, H2, 5-10m

Ptilocladia pulchra Sonder NE, 10m

Ceramiales - Ceramiaceae - Dasyphileae

Dasyphila preissii Sonder H2, 10m

Ceramiales - Ceramiaceae - Antithamnieae

Antithamnion hanovioides (Sonder) De Toni SE, NE, MO on Polvopes constricus, 0-10m

Ceramiales - Ceramiaceae - Spongoclonieae

Spongoclonium “sp. 3” NE, 10m

Spongoclonium sp. H2, 10m

Ceramiales - Ceramiaceae - Griffithsieae

Griffithsia monilis Harvey H1, 5m

Ceramiales - Ceramiaceae - Spyrideae

Spyridia dasyoides Sonder H1, W, 5-27m

Ceramiales - Ceramiaceae - Ceramieae

Ceramium pusillum Harvey H2, 10m

Ceramium rubrum C. Agardh H1, 10m

Dasyaceae

Heterosiphonia gunniana (Harvey) Reinbold NS, H1, 5-13m

Delesseriaceae-Nitophylloideae

Acrosorium ciliolatum (Harvey) Kylin H1, 5-11m

Apoglossum spathulatum (Sonder) Womersley & Shepley H1, 10m

Crassilingua marginifera (J. Agardh) Papenfuss SE, NW on Callophyllis, 10-23m

144 J. L. BAKER, G. J. EDGAR & N. S. BARRETT

Hymenena endiviaefolia (Hooker & Harvey) Womersley H1, H2, 5-10m

Hypoglossum revolutum (Harvey) J. Agardh H1, 10m

Laurencieae

Laurencia majuscula (Harvey) Lucas NE, 10m

Rhodomelaceae

Cliftonaea pectinata Harvey H2, 10m

Coeloclonium tasmanicum (Harvey) Womersley H2, 10m

Dictvomenia harveyana Sonder_ H1, H2, 5-10m

Epiglossum smithiae (Hooker & Harvey) Kutz. H1, H2, 5-10m

Herposiphonia versicolor (Hooker & Harvey) Reinbold H1, 10m

Laurencia brongniartii J. Agardh ~H1, 5m

Lenormandia pardalis J.Agardh H1, 5m

Protokuetzingia australasica Montagne H1, 5m

Transactions of the Royal Society of S. Aust. (2005), 129(2), 145-157.

INTERTIDAL MOLLUSCAN AND ECHINODERM DIVERSITY AT ALTHORPE

ISLAND AND INNES NATIONAL PARK, SOUTH AUSTRALIA

by K. BENKENDORFF*

Summary

BENKENDORFF, K, (2005). Intertidal molluscan and echinoderm diversity at Althorpe Island and Innes National

Park, South Australia. Trans. R. Soc. S. Aust. 129(2), 145-157, 30 November, 2005.

Species inventory data provide the first step towards understanding local marine communities. Here I apply

rapid biodiversity assessment to ten intertidal sites, with two levels of habitat complexity: rock platforms with

and without boulder fields. Five sites were located on Althorpe I., with a further five reefs in and around the

Innes National Park on Yorke Peninsula. One hour, timed-search surveys were used to determine the species

richness and rarity of macromolluscs and echinoderms. In total 82 molluscan species were found, but only eight

echinoderms. A large proportion of species was found to be numerically rare (<5 individuals per site) or spatially

rare (only observed at one or two sites). Reefs with boulder fields supported the highest species richness, and

overall more species were found on Althorpe I, (66) than the mainland (59). Multivariate analyses revealed that

different molluscan communities occur on rock platforms with and without significant boulder habitat. Different

molluscan communities also appear to occur on Althorpe |. when compared to the mainland sites, Thus, a

comprehensive marine reserve system should include representation from these two distinct intertidal rocky reef

habitats, at both offshore and coastal locations on Yorke Peninsula.

Key Worbs: Intertidal reefs; macromollusc; echinoderm; community composition; marine reserve selection;

species inventory.

Introduction

Species inventory data provide the foundation for

conservation and management of marine habitats.

Unfortunately, however, inadequate inventory data

are the “rule rather than the exception” for most

Australian marine habitats (Smith 2005). The

problem of access to many marine habitats means

that they are poorly known compared to their

terrestrial counterparts. As a consequence the

application of systematic reserve selection

procedures has been very limited in the marine

environment (Pressey & McNeill 1996; Gladstone

2002) and, instead, site selection for protective

measures has often been based on educated

guesswork (Smith 2005).

The limited resources available for marine species

inventories have led to the need for development of

rapid biodiversity assessment methods (see

Benkendorff & Davis 2002; Gladstone 2002;

Benkendorff 2003; Smith 2005). Typically, these

rapid assessment programs aim to establish whether

certain locations are representative and/or of

particular ecological importance to the region, within

a limited time frame. Species richness (a diversity) is

often used as an indicator of ecological importance

because it captures one important aspect of

biological diversity and can be reliably assessed with

"School of Biological Sciences, Flinders

GPO Box 2100, Adelaide, South

Email: kirsten.benkendorff@flinders.edu.au

University,

Australia S001.

relative ease (Benkendorff 2003). Species richness

inventories can be used to detect biodiverse hotspots

and facilitate the prioritization of sites for

conservation (Benkendorff & Davis 2002; Gladstone

2002), as well as produce estimates of the regional

(y) diversity (e.g. Warwick & Light 2002), and the

turnover of invertebrate assemblages between

locations (B diversity e.g. Oliver & Beattie 1996).

Species richness measures are nevertheless limited

to describing only one aspect of the potential

variation in ecological communities. Thus, many

ecologists prefer to collect abundance data to

calculate diversity indices (Bisby 1995), quantify

spatial patterns of intertidal diversity (e.g. Davidson

et al. 2004) or to undertake multivariate analyses of

the community composition (e.g. Underwood &

Chapman 1998). However, given the large natural

variation that is known to occur in invertebrate

communities on intertidal reefs, abundance data will

only be useful if sufficiently replicated in space and

time (e.g. Underwood & Chapman 1998).

Benkendorff (2003) found that the collection of

quantitative abundance data using replicated quadrats

can greatly reduce the area of reef that can be

sampled in limited time frames, thus negatively

affecting the measure of overall species richness. An

alternative approach used by Smith (2005) is to

provide a semi-quantitative ranking using defined

abundance categories for each species observed

during search surveys. This scoring system can be

used in multivariate comparisons of the species

composition across habitats or locations, whilst not

146 K. BENKENDORFF

adding greatly to the time taken to complete each

survey. Rarity in most marine species is poorly

understood (Chapman 1999) and thus any assessment

of numerical abundance and spatial distribution will

add to our current state of knowledge.

Rapid biodiversity assessment will typically

involve the use of one or more indicator taxa.

Previous studies on intertidal reefs in NSW, Australia

have demonstrated that macro-molluscs are effective

surrogates for intertidal diversity, with high

correlations with over-all species richness

(Gladstone 2002; Smith 2005). Molluscs also have

the advantage of being widespread and easily

sampled, with relatively well understood taxonomy

and ecology compared to other invertebrate taxa

(Benkendorff & Davis 2002, Smith 2005). The

species richness (a diversity) of intertidal reef

macromolluscs can be reliably assessed using short

(1 hr) unreplicated timed search surveys

(Benkendorff 2003), and these surveys have been

shown to produce accurate site rankings when

compared to longer-term species richness data

(Benkendorff & Davis 2002). Mollusc diversity can

vary substantially between intertidal reefs, with

habitat complexity likely to be a major driving factor.

In particular, species-rich hotspots on the NSW coast

are characterized by the presence of suitable boulders

for molluscan egg mass deposition (Benkendorff &

Davis 2004).

In southern Australian waters, 95% of marine

molluscs are estimated to be endemic (Allen 1999).

Overall, the southern temperate flora and fauna is

reported to have much higher levels of endemism

than those in the tropical north (Zann 2000), thus

highlighting the need for effective conservation in

this bioregion. However, most of the marine estate

under protection or management by the

Commonwealth Government is on the Great Barrier

Reef or off-shore e.g. in the Great Australian Bight

(Edyvane 1996; DEH 2004). Currently, only 0.26%

of the South Australian marine jurisdiction is under

protection in aquatic reserves (DEH 2003). In Gulf

St Vincent and Spencer Gulf, significant intertidal

reefs are protected at Aldinga, Troubridge Pt and

Whyalla. Notably, these are all limestone reefs with

no representation of granite or basalt reefs on the

S.A. mainland, Some near-shore islands with granite

reefs, such as West I. and Goose I. are included in

aquatic reserves. These islands may provide

intertidal habitat that is relatively less impacted by

humans than the mainland. Thus, an additional

reserve around the Althorpe Is could prove beneficial

for the conservation of local marine biodiversity. A

' Gaut, A, (2000). A preliminary amateur intertidal study of

Althorpe Island Conservation Park. Friends of the Althorpe

Islands Conservation Park, Unpublished report.

comparative investigation of the community

structure is needed to determine whether both

offshore and near-shore reefs, encompassing the full

range of substrata found in the region, should be

represented in future marine conservation areas.

Only one unpublished study has previously

investigated the biodiversity of intertidal reefs on

Althorpe I'. Molluscs were found to be the most

numerous group, with a total of 17 species identified

and shell evidence for several more. This list is likely

to be an underestimation of biodiversity due to high

swell conditions and the broad scope of the survey.

Consequently, further assessment of the intertidal

biodiversity is warranted.

The main objective of this study was to use rapid

biodiversity assessment surveys to obtain species

inventories of the macromollusc and echinoderm

fauna occurring at representative rocky intertidal

sites on Althorpe I. and around Innes National Park

on Yorke Peninsula mainland. The numerical and

spatial rarity of each species in this region was also

investigated. Multivariate analyses on the molluscs

were then used to compare the community

composition between the mainland and island

locations. The influence of habitat was also assessed

by comparing simple rock platforms to reefs with

significant boulder fields across both locations. This

study should help inform planning for a

comprehensive and representative system of marine

reserves in South Australia.

Methods

This study was undertaken at five intertidal sites

along the rocky coast of Althorpe I., with a further

five sites on the mainland of Yorke Peninsula, South

Australia (Fig. 1). The mainland sites are collectively

referred to as Innes National Park, although only four

sites actually occur in the Park, with the remaining

one ~10 km north at Marion Bay. The surveys were

all conducted within the same week (6th — 12th

February, 2004) over low tide (0.12 — 0.16 m, Table

1). At both the island and mainland locations, sites

were selected according to the accessibility of habitat

in the lower littoral zone and thus all sites are

relatively gently sloping with some protection from

the prevailing SW swell. All sites encompass areas of

rock platform with rock pools and crevices. At each

location, three sites were complex granite rock

platforms with mixed granite, basalt and limestone

boulder fields encompassing >10% of the total

search area (Table 1). The remaining two sites at each

location were predominantly rock platforms with

<1% boulder habitat. These reefs were composed of

granite, with the exception of one mainland site,

Stenhouse Bay, which is a limestone reef (Table 1).

Comparable limestone reefs do not occur on

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A.

147

YORKE

PENINSULA

{

Gym Beach

eenuneee

INNES “Sennen,

NATIONAL *

PARK Marion Bay

Stenhouse Bay

Cape Spencer 5 ALTHORPE I.

INVESTIGATOR

STRAIGHT

NOT TO SCALE

Fig. 1. Intertidal study sites on Althorpe Island and Innes National Park, Yorke Peninsula, South Australia.

148 Kk. BENKENDORFF

TABLE |. Description of the intertidal sites surveyed for molluscs and echinoderms on: a) Althorpe I.; and b) Innes National

Park, Yorke Peninsula, South Australia.

Althorpe I. Site GPS Reading Tide Habitat

1) Western Channel S 35° 22.081" 0.15 Granite channel filled with basalt and some limestone boulders

E 136° 51.333’

2) Western Platform S 35° 22.046’ 0.15 Granite rock platform

E 136° 51.352’

3) The Fangs S 35° 22.195" 0.14 Granite rock platform with deep pools

E 136° 51.999’

4) East of Mooring Bay S 35° 22.165’ 0.14 Granite rock platform extending into a shallow basalt, granite

E 136° 51.793’ and limestone mixed boulder field

5) West of Pareora S 35° 21.989’ 0,12 Granite platform with channels of granite, basalt and

Monument E 136° 51.603’ limestone boulders

Mainland Site GPS Reading Tide Description

1) Gym Beach S 35° 09.230" 0,12 Granite rock platform and boulder field

E 136° 54.445’

2) Marion Bay South —S 35° 14.978" 0.12 Granite platform with some deep pools

E 136° 58.937’

3) Cape Spencer SWS 35° 17.954" 0.15 Basalt and granite boulders extending into granite platform

E 136° 52.811"

S 35° 17.932" 0.15

E 136° 53.016’

S 35° 16.025’

E 136° 57.250'

4) Cape Spencer NE

5) Stenhouse Bay

Mixed granite, limestone and basalt boulder-filled inlet

with granite outcrop and platform

Limestone platform

Althorpe I., although they are common on the coast

around Innes National Park; consequently Stenhouse

Bay was included to provide a more comprehensive

survey of the regional biodiversity.

Using measuring tapes, a 20 x 20 m quadrat was

defined as the search area, commencing from the

lowest possible level of the shore at each site. A GPS

reading was taken from the middle of the site (Table

1). One hour timed search surveys were then

conducted at each site according to Benkendorff

(2003), with the modification that the search was

divided into 6 x 10 min time intervals to enable the

construction of species accumulation curves. The

surveys involved searching for and recording all

molluscan and echinoderm species (>5 mm) found

within the entire range of intertidal habitats present

within the 20 x 20 m plot. Molluscs were identified

according to Lamprell & Healy (1998), Lamprell &

Whitehead (1992), Wilson (1993), Edgar (1997),

Jansen (2000), Coleman (2003) and Rudman (2004).

Voucher specimens were collected from species that

could not be easily identified and some of these were

later identified with the assistance of Peter Hunt (SA

Malacological Society). Higher taxonomic

groupings of the Mollusca have been assigned

according to the classification outlined in Beesley et

al. (1998). Echinoderms were identified according to

Edgar (1997).

Two measures of species rarity have been assessed

in this study: 1) spatial rarity; and 2) numerical rarity.

Spatial rarity is simply defined by the frequency of

sites at which each species was recorded. Numerical

rarity was assigned according to maximum local

abundance of each species observed at each site in an

hour. Species were assigned to one of the following

categories: a) rare, <5; b) uncommon, 5 — 20; c)

common, 21 — 100, and; d) abundant, >100

individuals. Frequency distributions for numerical

rarity are calculated for the region by taking the

maximum abundance for each species at any one

site.

Multivariate analyses were undertaken to

investigate the molluscan communities using the

Primer Software package Version 5 (Clarke & Gorely

2001). Using semi-quantitative abundance scores,

the similarities between each pair of sites were

analysed using the Bray-Curtis measure according to

Smith (2004). Non-metric multi-dimensional scaling

(nMDS) was performed on the similarity matrix and

portrayed in two-dimensional plots using the

grouping factors of: a) location (Althorpe I. vs.

mainland) and b) habitat (rock platforms vs. platform

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A. 149

TABLE 2. Species list for a) Mollusca and b) Echinodermata, at ten intertidal rocky reef sites on Athorpe I. and in or near

Innes National Park, Yorke Peninsula. * = egg masses were also observed. No. sites refers to the number of sites each

species was recorded from within each location (max = 5). The relative rarity of each is defined according to the

maximum number of individuals observed at any one site within the location: Rare (R) <5; Uncommon (U), 5-20;

Common (C), 21 — 100; Abundant (A), >100 individuals.

Family Species Althorpe Island Innes National Park

No. sites Rarity No. sites Rarity

Eogastropoda

Patellidae Scutellastra chapmani

Patella peronii

Nacellidae Cellana tramoserica

Cellana solida

Lottidae Notoacmea petterdi

Notoacmea mayi

Patelloida cf. flammea

Patelloida alticostata

Patelloida latistrigata

AMANNOMNBNN

AMaAWaAnnNnN ©

Orthogastropoda

Neritopsidae Nerita atramentosa

Haliotidae Haliotis rubra

Fisurellidae Scutus antipodes

Clypidina rugosa

Notomella candida

QACAAASF ANAANAPPPrACA

Trochidae Austrocochlea rudis

Austrocochlea adelaidae

Austrochoclea odontis

Notogibbula preissiana

Clanculus brunneus

Granata imbricata

Talobis rotunda

Canthrindella picturata

Herpetoma pumilo

ARRAYS!

CAAA QPS SSF COACH SFPAYPLrSrIrA:

Thalotia c.f. chlorostoma

Talopena c.f. gloriola

Turbinidae Turbo undulatus

Turbo torquatus

Micrastaea aurea

Qn:

ZFaQ'A

Austroliotia botanica

Subninella gruneri

Astralium aureum

Cererthidae Bittium granarium

Littorinidae Bembicium nanum

Littorina acutispira

Littorina unifasciata

Nodilittorina praetermissa

Rissoina crassa

Rissoina fasciata

Calyptraeidae Crepidula aculeate

Epitoniidae Opalia australis

Ranellidae Sassia sp.

Cancellariidae Cancellaria c.f. granosa

Buccinidae Cominella lineolata

Cominella eburnea

Unidentified sp. 1

Columbellidae Mitrella pulla

FB OP OCOKF NON COWWHENWNNWN | HN

ANF SSC:

AAAAAQVPSPePCA!

1 wi

KEEP ENON HKFNHKHNUUNUNWWHK OH HOW SCH OH KE UNSCANHK HK HY

An!

SOF ON OCHONUUN

AACLY!

150

TABLE 2. Cont.

K. BENKENDORFF

Family Species Althorpe Island Innes National Park

No. sites Rarity No. sites Rarity

Muricidae Lepsiella vinosa 2 i, 2 €

Dicathais orbita 5 Cc 5 Cc

Lepsiella flindersii 1 R 1 R

Marginellidae Austroginella sp. 1 R 0 -

Conidae Conus anemone 2 R 3 R

Mitridae Mitra badia 2 R 0 -

Cerithiopsidae Euselia pileata 1 R 0 -

Unidentifed Unidentified sp. 2 1 R 0 -

Siphonartidae Siphonaria funiculata 5 C I U

Siphonaria zelandica 2 U 2* €

Siphonaria diemenensis aie A 4 A

Siphonaria c.f. baconi 4 Cc 4 A

Anapsidae Aplysia parvula ii R 1 R

Notospidae Berthellina citrina 1 R 0 -

Unidentified sp. 3 0 - 1 R

Nudibranchia Austreolis ornata 1 R 0

Rostangia bassia 1* R 0 -

Unidentified eggs 0 - 1* R

Flabellina poenicia 0 - 1 R

Polyplacophora

Ischnochitonidae Ischnochiton elongatus 5 C 3 C

Ischnochiton variegatus 5 C 2 U

Ischnochiton australis 0 - 2 R

Mopalidae Ischnochiton sp. 1 R 0 a

Plaxiphora albida 3 U 4 Cc

Bivalvia

Galeommatidae Lasea australis 5 € 3 C

Carditidae Cardita sp. 3 R 1 R

Ostreidae Ostrea angasi 0 - 1 U

Saccostrea glomerata 1 R 0 -

Mytilidae Hormomya erosa 0 - 1 R

Xenostrobus pulex 1 C + A

Venerupis c.f. anomala 2, R 2; R

Pteriidae Pinctada sp. 1 R 0 -

Archidae Barbatia riculata 0 - 1 R

Teredinidae Thraciopsis subrecta 1 R 0 -

Unidentified Unidentified sp. 4 0 - 1 R

Cephalopoda

Spirulidae Spirula spirula 0 - 1 R

Family Species Althorpe Island Innes National Park

No. sites Rarity No. sites Rarity

Asteroidea

Asterinidae Patiriella calcar 2 R 2 U

Asteriidae Coscinasterias muricata 1 R 0 -

Goniasteridae Tosia australis 1 R 0 -

Oreasteridae Nectria sp. 1 R 0 -

Ophiuroidea

Ophionereididae Ophionereis schayeri 3 R 2; R

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A. 151

TABLE 2. Cont.

Family Species Althorpe Island Innes National Park

No. sites Rarity No. sites Rarity

Ophiactidae Ophiactis resiliens 1 R 0 =

Echinoidea

Temnopleuridae Holopneustes inflatus 1 R 0 7

Echinometridae Heliocidaris erythrogramma l R 0 7

No. Species

0 10 20 30

Time (mins)

= Western Channel

@- Western Platform

’- The Fangs

- East of Mooring Bay

@- West of Monument

—O— Gym Beach

——*— Marion Bay

—*— SW Cape Spencer

-——e— NW Cape Spencer

—*— Stenhouse Ba’

40 50 60

Fig. 2. Species accumulation curves for molluscs recorded during one-hour search surveys at five intertidal sites on

Althorpe Island (broken lines) and five intertidal reefs on the Yorke Peninsula mainland (Innes National Park, solid lines).

with boulder fields). Significant differences in

community structure were then explored using a two-

way ANOSIM. SIMPER was undertaken to identify

the species contributing to the differences in

community structure. These analyses were repeated

using a presence/absence transformation on the

species matrix.

Results

A total of 82 species of molluscs and eight

echinoderms was recorded during the 10 surveys on

Yorke Peninsula and Althorpe I. (Table 2). Sixty-six

of the molluscan species were observed on Althorpe

I., including 23 species that were not found on the

mainland (Table 2a). Fifty-nine species of molluscs

were recorded in Innes National Park, with 15 not

observed on Althorpe I. The egg masses of five

gastropods were observed on Althorpe I., whereas

the egg masses from four species were observed on

the mainland (Table 2). Eight species of

Echinodermata were recorded and all of these were

found on Althorpe I., including six species that were

not observed on the mainland (Table 2b).

The maximum number of species recorded at any

one site was 46 molluscs (Fig. 2) and three

echinoderms in the Western Channel at Althorpe I.

Nearly half this number of species was recorded at

Stenhouse Bay in Innes National Park, with 25

molluscs (Fig. 2) and no echinoderms. Species

152 K. BENKENDORFF

25 @ Molluscs

O Echinoderms

a) 2

xe)

1 2 3 4 5 6 7 8 9 10

Number of sites

no 30

b) f %

ce!

wae 15

is)

0 T T 1

rare uncommon common abundant

Fig. 3. The frequency distribution of rarity in molluscs and echinoderms recorded during ten intertidal surveys on Althorpe

Island and at Innes National Park; a) spatial rarity as determined by the number of sites occupied by each species; b)

numerical rarity, as determined by the maximum abundance at any site: Rare (R) < 5; Uncommon (U), 5-20; Common

(C), 21-100; Abundant (A), >100 individuals.

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A. 153

Stress: 0.0

a Althorpes

y_ Innes National Park

A. Mixed boulders

v_ Granite platform

Limestone platform

Fig. 4. Two-dimensional nMDS plot for the molluscan community data based on semi-quantitative abundance rankings

from 10 intertidal sites. The plots show dissimilarities between sites grouped according to: a) location (Althorpe I. vs.

mainland); and b) habitat (rock platforms without boulders vs. mixed boulders with some rock platform).

richness accumulation curves show that new species

were still being recorded in the last 10 minutes of the

survey at most sites (Fig. 2). In particular, the species

accumulation curves were still rising steeply for the

two most species-rich sites, the western channel on

Althorpe I and Gym Beach in Innes National Park.

Overall, there was no apparent difference in

cumulative species richness for sites on Althorpe I.

when compared to the mainland (Fig. 2). However,

the four sites which are relatively simple rock

platforms have the lowest accumulated species

richness compared to those sites with significant

boulder habitat (Fig. 2, Table 1).

The majority of species recorded in these surveys

was found to be spatially and/or numerically rare

(Fig. 3). Thirty four percent of molluscs and 75% of

echinoderm species were only detected at a single

site (Fig. 3). Furthermore, a maximum of only one or

two individuals was recorded for 49% of mollusc and

87.5% of echinoderm species (Fig. 3b). No

echinoderm species were recorded at more than half

of the sites or were numerically common at any one

site (Fig. 3a). By comparison, 10 species of molluscs

were recorded at all sites and these species, along

with an additional four were observed to be

numerically abundant (Fig. 3a), with 100s of

individuals at most or all sites.

The nMDS ordination of the full data set using

semi-quantitative abundance categories indicates

that discrete molluscan assemblages occur on

Althorpe I. compared to those on the mainland

(Innes National Park, Fig. 4a). Furthermore,

segregation of the sites can be observed according to

the molluscan communities occurring on the

different types of intertidal habitats (Fig. 4b, stress

0.07). The sites with mixed boulder fields are

154 kK. BENKENDORFF

clustered reasonably tightly compared to the rock

platforms. The limestone platform at Stenhouse Bay

is somewhat separated from the granite reefs. Two-

way ANOSIM reveals significant differences in the

community composition between rock platforms and

boulder reefs (Global R = 0.542, p = 0.03), but

differences between the island and mainland

locations are only significant at the 10% level

(Global R = 0.514, p = 0.067). ANOSIM using

presence/absence transformed data supports the

above trends with significant differences according

to habitat (R = 0.604, p = 0.03), but not location (R

= 0.4, p = 0.133).

SIMPER analyses using the abundance matrix

indicate that the average dissimilarity between

Althorpe I. and the mainland locations is 37.67% of

the species composition. Cumulative contributions

from 51 species are required to account for over 90%

of the total dissimilarity between locations; 30 of

these species have dissimilarity/standard deviation

ratios above or close to 1, and are therefore

considered reliable indicators. A subset of these

species that contribute to >3% of the average

dissimilarity are presented in Table 3a. SIMPER

analyses on the presence/absence transformed data

also revealed Siphonaria funiculata and Xenostrobus

pulex to be reliable indicators, with the former more

prevalent on Althorpe I. and the latter more prevalent

on the mainland.

Investigation of the dissimilarity between rock

platform and mixed boulder communities using

SIMPER on the abundance matrix revealed an

average dissimilarity of 37.8%. Again, 51 species

contribute to >90% of the total dissimilarity and 35

have dissimilarity/standard deviation ratios above or

close to one. The subset of species contributing to

over 3% of the dissimilarity is listed in Table 3b. In

particular, Austrocochlea spp. appear to be useful

indicators for the different habitats, with A.

concamerata and A. odontis more abundant in

boulder communities, whereas A. adelaidae is more

prevalent on rock platforms. Presence/absence

transformation supports use of A. concamerata as an

indicator for molluscan communities in boulder

habitats.

Discussion

A relatively high species richness of molluscs, but

low richness of echinoderms can be found on rocky

intertidal reefs around Althorpe I. and Innes National

Park. This study has contributed an additional eight

echinoderm species and 50 molluscan species to the

preliminary study of the Althorpe intertidal

communities conducted by Gaut'. Species lists are an

important first step towards conservation and

management (Oliver & Beattie 1993; Soule & Kohm

1989; Stork 1994), because they facilitate the

identification of rare species and enable quantitative

measures of biodiversity. However, comprehensive

species lists are very difficult to obtain due to

temporal variation in marine communities. So this

study should only be regarded as a representative

snapshot of the local fauna.

It can be difficult to compare the results of marine

biodiversity surveys between studies and across

regions due to lack of standardization in the survey

methods (Benkendorff 2003). Nevertheless, the

mean species richness recorded per site for molluscs

in this study (31.7+6.4) is comparable to previous

timed search surveys on the NSW Coast (e.g.

Gladstone 2002, mean = 31+8.9 from three-hour

searches; Benkendorff 2003, mean = 38.9+12.2 from

one-hour searches). The total number of molluscs

(y diversity) recorded on Yorke Peninsula during this

study (81 from 10 hours searching, Table 2a) also

compares favorably to that recorded from intertidal

reefs along the south east coast of Australia. For

example, Gladstone (2002) reports a total of 80

molluscan species from a total of 90 search-hours at

15 locations along the NSW Coast, and Smith (2005)

reports 170 molluscs from 25.8 total search hours on

eight headlands around the Solitary Is National Park,

NSW. In the same study by Smith (2005), 30

echinoderm species were recorded, indicating the

relatively low diversity for this Phylum on intertidal

reefs around Yorke Peninsula, S.A. (eight species,

Table 2b).

Across both island and mainland locations, sites

with boulder habitat were found to have higher

species richness than the simpler rock platforms

(Fig. 2, Table 1). This finding is consistent with the

Habitat Diversity Hypothesis (Connor & McCoy

1979), which predicts that a greater diversity of

species will occur where a greater diversity of

habitats exists. A boulder-filled channel on the NW

side of Althorpe I. (Fig. 1) was found to be

particularly rich in intertidal molluscs (Fig. 2).

According to Benkendorff & Davis (2002), this site

could be regarded as a regional “hotspot” of

molluscan diversity by applying the definition of two

standard deviations above the regional mean. No

other sites were found to fit this “hotspot” definition,

although Gym Beach in Innes National Park is close

(Fig. 2). The continuously rising species

accumulation curves at both these sites suggest that

species richness may be underrepresented by the

one-hour search surveys used. Benkendorff & Davis

(2002) have demonstrated that it is significantly

more difficult to obtain a comprehensive species list

at species-rich intertidal hotspots. Furthermore,

relatively few egg masses were recorded at any site

during this study, most likely due to the timing of the

surveys (late summer). However, both Gym Beach

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A. 155

TABLE 3. SEMPER results for the difference in molluscan communities between: a) Alihorpe I. and mainland (Innes NP)

locations, and b) rock platforms and mixed boulder reefs. The Bray Curtis dissimilarity measure is based on a categorized

rarity matrix with 999 permutations and several repeated computations. The average abundance is taken from semi-

quantitative abundance rankings averaged across each group of sites for each species, such that 0 = not present; 1 = rare

(<5 individuals); 2. = uncommon (5-20 individuals), 3 = Common (21-100 individuals); 4 = Abundant (>100

individuals). Only those species with a dissimilarity/standard deviation ratio close to and above one are listed as

consistent indicators and only those contributing to greater than 3% of the total dissimilarity are presented here.

Species Average Abundance Ratio % Cumulative

Althorpe I. Innes NP Diss/SD contribution — Contribution

Siphonaria funiculata 3 > 0.4 2.86 4.66 4.66

Xenostrobus pulex 0.6 < 2.8 1.47 4.28 8.94

Turbo undulatus 1.2 < 2.8 1.4 3.12 12.05

Bembicium nanum 2 < 2.1 1.36 3.08 15.14

Plaxiphora albida 0.8 < oy 1.66 3.04 18.18

Austrocochlea odontis | < 1.6 0.98 3,02 21.20

Species Average Abundance Ratio % Cumulative

Mixed boulders Rock platforms Diss/SD = -% contribution Contribution

Austrocochlea concamerata 3 > 0.25 3.59 4.9] 4.91

Austrocochlea adelaidae 0 < 2 0.97 3.51 8.42

Austrocochlea odontis 2 > 0.25 L.11 3.38 11.79

Bembicium nanum 2.5 > 1.25 1.43 3.35 15.14

Xenostrobus pulex 1.67 < 1.75 1.07 3.29 18.43

Notogibbula lehmanni 2.83 > I 1.52 3.22 21.65

and the Althorpe channel provide extensive boulder

habitat that would be suitable for egg mass

deposition by a wide range of gastropods (pers. obs.,

see also Benkendorff & Davis 2004). Consequently,

additional surveys at these sites during late spring

when more molluscs are breeding (pers. obs.) may

further highlight the importance of these sites as

habitat for intertidal molluscs.

Species richness, community composition and

rarity should all be taken into consideration for

biodiversity conservation and management. Species

turnover (B diversity) was high between sites in both

locations, with 28 molluscs and six echinoderms

only recorded at a single site (Fig. 3a). This suggests

that any one hotspot will not fully represent all the

spatially rare species observed in this study.

However, a new marine conservation reserve around

the Althorpe Is would encompass all the associated

intertidal reefs surveyed here. All but 15 molluscan

species that were recorded solely on the mainland in

this study would be represented in such a reserve

(Table 2). Overall, a higher species richness and

more rare species were recorded on Althorpe I. when

compared to the mainland (Table 2). Whilst this

supports prioritization of Althorpe I. for protection in

anew aquatic reserve, this location should still not be

considered representative of the mainland. Some

distinct clustering could be seen for the Island versus

the mainland sites in the nMDS ordination plot (Fig.

4a) and ANOSIM indicated a marginally significant

difference in the community composition between

these two locations. A large number of species were

found to contribute to the average dissimilarity

between locations, with many species such as

Xenostrobus pulex being more common on the

mainland. However some, such as Siphonaria

Juniculata, are more prevalent on Althorpe I. (Table

3a). The reasons for these differences in molluscan

community structure between Althorpe I. and Innes

National Park are currently unclear but are likely to

be due to a combination of biotic and abiotic factors.

In particular, distance from source populations and

current patterns would influence the chance of

populations establishing at the various sites.

In addition to representation of the coastal and

mainland locations, there is also a clear need to

protect the different types of intertidal habitats in

marine reserves. Distinct molluscan communities

were found on rock platforms, compared to those

sites with mixed boulder fields, as shown by nMDS

ordination (Fig. 4b). Indeed, ANOSIM revealed that

the difference between habitats was more significant

156 Kk. BENKENDORFF

than that between the mainland and island locations.

The sites are separated for habitat type along a

different axis to that for location (Fig. 4), suggesting

that different molluscan species are driving these two

patterns, as supported by SIMPER analysis (Table 3).

Again, a large number of species were found to

contribute to the dissimilarity between habitat types,

and whilst many of these species are specialized to

live under boulders (data not shown), this does not

apply to those species with the greatest percent

contribution (Table 3b). Several Austrocochlea spp.

were identified as possible indicators for the

different communities, with A, adelaidae only

recorded on rock platforms without boulders (Table

3b).

Not all of the variation in molluscan communities

can be explained by the two factors used in this study.

Notably, there was a fairly high degree of separation

between all sites in the nMDS ordination (Fig. 4). In

particular, there is an apparent separation between

the granite platforms compared to the limestone reef

at Stenhouse Bay (Fig. 4b). However, the lack of

replicate sites prevents any further conclusions being

drawn about the effects of substratum type on

intertidal molluscan community composition. At the

mixed boulder sites, the total area covered by

boulders and the proportion of different rock types

may account for some more of the variation. Thus,

the effects of substratum and other abiotic factors,

such as wave exposure, are worthy of further

investigation.

The patterns in species richness and community

composition observed in this study appear to be

largely driven by rare species (Fig. 3). Rare species

are usually regarded as those that persist with

naturally low abundances and/or restricted

geographic ranges (Gaston 1994; Chapman 1999).

The species abundance distribution (numerical

rarity) for intertidal molluscs and echinoderms in

this study is skewed to the right (Fig. 3b), indicating

that the majority of species are found in very low

numbers and most of the individuals belong to a few

abundant species. This pattern is consistent with

previous studies on a wide range of communities

(e.g. Fisher et al. 1943; MacArthur 1957). The

frequency distribution for the spatial occurrence of

intertidal molluscs in this study is also skewed right,

but tending toward bimodality (Fig. 3a). This

indicates that there is one group of molluscs whose

presence is predictable, on most, if not all intertidal

reefs in this S.A. bioregion, whilst most other species

are highly unpredictable with patchy or localised

distributions. Bimodal distributions for the number

of locations that intertidal species are recorded have

been previously observed at the regional scale (e.g.

Gladstone 2002). Gaston (1994) suggests that these

distributions could, in part, reflect sampling artifacts

because the left hand mode is likely to be inflated

due to the fact that species with low abundances are

harder to find and therefore have a lower probability

of being recorded at any site. In support of this, the

majority of numerically rare species were only found

at one or two sites in this study (Table 2). Species

that are both numerically and spatially rare are often

termed accidentals, vagrants, immigrants or

incidentals (Gaston 1994) because they do not

appear to maintain viable populations in the study

area. Indeed, the interconnectedness of intertidal and

subtidal habitats could create problems for assessing

the rarity of intertidal species due to vertical

migration. Nevertheless, from an_ ecological

perspective, immigrants should be regarded as a part

of the community because they could contribute to

species interactions (Gaston 1994). The potential

conservation importance of these rare species should

not be overlooked until more is known about their

source populations.

In conclusion, this study demonstrates that the

intertidal reefs around Althorpe I. and Innes National

Park support a high species richness of molluscan

fauna, as well as a high proportion of apparently rare

molluscan and echinoderm species. Due to

differences in the community composition, a

comprehensive marine reserve system should

include both island and mainland sites. Furthermore,

representative sites need to encompass the full range

of different intertidal habitats that occur in the

region, at least including boulder fields and rock

platforms. The importance of geology should be

further investigated since only limestone reefs are

currently protected on Yorke Peninsula mainland.

Granite reefs occur on both Althorpe I. and in Innes

National Park. These are often accompanied by

mixed basalt, granite and limestone boulder fields

providing high habitat complexity and greater

species richness compared to the rock platforms.

One such site on Althorpe I. was identified as a

hotspot of species richness that could be prioritised

for conservation. But ultimately, the large number of

spatially rare species found in the region and the

large variability in community composition between

sites indicates the need to protect a suite of sites

rather than any one location.

Acknowledgements

1 am extremely grateful to Alex Gaut for her

assistance in the field. This research would not have

been possible without the support of R. K. Lewis,

and the help of Sue Murray-Jones and Scoresby

Shepherd for organizing this scientific expedition to

Althorpe Is, including access to the island on the

SARDI research vessel Ngerin. Thanks also to the

Friends of the Althorpes for providing

INTERTIDAL COMMUNITIES OF ALTHORPE I. AND MAINLAND, S.A. 157

accommodation on the Island and for their interest

conserving the local environment. I am also grateful

to the Rangers in Innes National Park for allowing

access and collection permits for the intertidal reefs

within the Park. The assistance of Peter Hunt in the

identification of some smaller molluscan species is

much appreciated. Constructive comments from two

anonymous reviewers have improved the manuscript.

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Transactions of the Royal Society of S. Aust. (2004), 129(2), 158-169.

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

by D. A. STAPLES*

Summary

StapLes, D. A. (2005). Pycnogonida of the Althorpe Islands, South Australia. Trans. R. Soc, S. Aust. 129(2),

158-169, 30 November, 2005.

Six species of shallow-water Pycnogonida are recorded from the Althorpe Islands. Pseudopallene watsonae sp.

nov. and Pseudopallene inflata sp. nov. are described. The adult male of Spasmopallene reflexa (Stock, 1968),

described on the basis of a juvenile, is described and the species rediagnosed. The genera Spasmopallene Stock,

1968 and Pallenella Schimkewitsch, 1909 are synonymised with Pseudopallene Wilson, 1878. The holotype of

Ps. ambigua Stock, 1956 has been re-examined and compared with the new species. Callipallene emaciata

micracantha Stock, 1954 is raised to species status. Stlopallene cheilorhynchus Clark, 1963 and Achelia

transfugoides Stock, 1973 are recorded.

Key Worps: Pycnogonids; Althorpe Islands; Callipallenidae; Ammotheidae.

Introduction

The pycnogonid fauna of South Australia is well

represented in museum and private collections

principally as a result of research and monitoring

programs conducted over many years. Most

specimens have been collected in Spencer Gulf and

Gulf St Vincent, with few records from offshore

islands, and none from the Althorpe Islands. Six

species referable to four genera and to two families

are represented. This collection provides an

opportunity to reconsider the generic status of

Spasmopallene and Pallenella and describe new

species. Specimens were collected using SCUBA at

depths ranging from 2 to 27 m. Background

information to the Althorpe Is and expedition details

are given by Murray-Jones & Shepherd (2005).

Specimens examined are lodged in the South

Australian Museum (SAM) and Museum Victoria

(NMV).

CALLIPALLENIDAE Hilton, 1942

Pseudopallene Wilson, 1878

Pseudopallene Wilson, 1878: 202; Stock, 1954.

Pallenella Schimkewitsch, 1909b: 1-13

(new synonomy).

Spasmopallene Stock, 1968: 39-40

(new synonomy).

Type species

Pseudopallene circularis (Goodsir, 1842).

Diagnosis

Scapes l-segmented, chelae fingers smooth;

proboscis conical distally, tip mamilliform; ovigers

“Museum Victoria, GPO Box 666. Melbourne, Victoria 3001.

Email: dstaples@museum.vic.gov.au

10-segmented, sexually dimorphic, terminal claw

well developed; palps and auxiliary claws absent. All

Australian species recorded to date share the same

habitus as the holotype; body compact, segmented;

anterior region of cephalon bulbous; neck short;

proboscis directed ventrally, not visible in dorsal

view. Scapes carried ventrally and angled outwards,

chelae directed inwards such that the fingers lay

directly below the mouth; cephalon and chelifores in

combination, assuming a somewhat triangular aspect

when viewed anteriorly. Juveniles and subadults

differ from adults in the shape of the proboscis and

chelifores, which are angled towards posterior;

chelae geniculate, directed forward and upward;

movable fingers distorted, digitiform; propodal heel

more pronounced; coxa 2 short; oviger spine

numbers low.

Remarks

The genus is common in southern Australian

waters. Species determination is confused by a range

of intermediate and overlapping characters, which no

doubt contributed to Clark’s (1963: 33) observation

that no two males (or females) of this group have

been found to be exactly alike. To a large extent this

variability can be attributed to ontogenetic changes

previously noted by Stock (1973a: 116) and Staples

(1997: 1053). In the present material, limited

variation is also evident in adult specimens,

particularly in the heel and oviger spine numbers.

Stock (1968: 39) erected the genus Spasmopallene

to accommodate specimens from the Great

Australian Bight, which he described as S. reflexa

and S. clarki Stock, 1968, at the same time

embracing Pseudopallene dubia Clark, 1963. It is

evident by association of larvae, juveniles and

subadults with adult Pseudopallene and the sharing

of a common substrate, that the genus

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

159

2.00MM

Fig. 1. Pseudopallene ambigua Stock, 1956, holotype: A, cephalon ventro-anterior; B, ocular tubercle; C, oviger claw;

D, trunk, lateral; E, third leg; F, trunk dorsal view; G, trunk, anterior view; H, propodus and tarsus, leg 3; I, chela.

Spasmopallene is based on juvenile Pseudopallene.

Sexual dimorphism of the ovigers is clearly evident

in subadults, perhaps contributing to an assumption

that the type specimens of Spasmopallene were

sexually mature. Neither author mentioned the

presence of genital pores in type material.

The genus Pallenella was erected by

Schimkewitsch (1909) to accommodate Pallene

laevis Hoek, 1881, principally based on that species

having 2-segmented scapes. In all other respects,

diagnosis of the genus agrees with Pseudopallene.

Hoek’s description of the scapes is, however, rather

ambiguous and appears to have been misinterpreted

by Schimkewitsch. His figures, which show the

scapes to be clearly segmented (PI.X1 Figs 8 and 9),

are inconsistent with his description of the scapes as

being “constricted at the base, and indistinctly

divided into two joints”. Examination of specimens

in this collection, and from elsewhere in southern

Australia, shows that a basal constriction of the scape

is a variable character present in more than one

species of Pseudopallene. In some instances the

restriction is well defined and with a suggestion of a

suture, but never segmented. It can be present in

160 D. A. STAPLES

subadult and adult specimens and, in some cases, in

specimens from the same collection. Pallenella has

remained monotypic and retention of the genus is not

supported by these observations. The genera

Spasmopallene and Pallenella are synonymised with

Pseudopallene.

Key to adults of species of southern Australian

Pseudopallene

1. a) Long segments of legs each with 2 distinct

annular constrictions ............... P. pachychiera

b) Leg segments without distinct constrictions .2

2. a) Propodal heel with spines arranged in 4 pairs

P reflexa

b) Propodal heel spines not paired...............0 3

3. a) Anterior margin of cephalon evenly rounded4

b Anterior margin of cephalon with mid-dorsal

MOU sscazsascesepeceesnenyedees itp svednnenreesreanacoaseionaeee 5

4. a) Terminal oviger claw, smooth, crenulate or

serrated on 1 margin only .........ceceeeeeeseeeeeee 5

b) Terminal claw serrated on both margins....... 6

5. a) Terminal oviger claw smooth or crenulate......

J cbbeaantn nested dobetaleg th abeiiaga renege eeetezssaaae P. ambigua

b) Terminal oviger claw serrated............ P. laevis

6. a)Chela fingers about '/+ length of palm,

immovable finger strongly curved, tip blunt...

ohh ES asta Sek So de at RENE aE P. watsonae

b) Chela fingers longer than '/2 length of palm,

immoveable finger weakly curved or

straight, pointed .......e cc eeeeeeeeeeeeeees P. inflata

Pseudopallene ambigua Stock, 1956

Fig | A-I

Pseudopallene ambigua Stock, 1956: 40-42,

Fig. 5 a-i; Clark, 1963: 31-33, Fig. 16 A-F;

-Stock, 1973a: 115-117, Fig. 7 c, g, e.

Material examined

Holotype, male. Bass Strait, Museum Hamburg,

K17 680.

Remarks

The holotype of Pseudopallene ambigua has been

re-examined for comparison with the new species.

The following observations can be made but

generally there is little to add to the original

description. Stock remarked on the unusual shape of

the ocular tubercle, which he illustrated with a

stepped anterior margin (Stock, 1956: Fig. 5). No

such unusual feature is now evident, suggesting

perhaps, temporary handling damage to the tubercle.

The scapes have a slight basal constriction and coxa

2 is more inflated distally than figured. Re-

examination of the oviger terminal claw using high

resolution digital imaging shows that the striations

observed by Stock are in fact crenulations (Fig. 1, C).

Pseudopallene ambigua shares much in common

with P /aevis. Both species are of similar size (leg

spans > 43 mm and about twice the size of the new

species described herein) and share Bass Strait as

their type locality. In both type specimens, only the

inner margin of the oviger terminal claw is serrated

or bears striations, whereas all specimens in this

collection bear distinct serrations on both margins. I

am grateful to Dr. H. Dastych of the Hamburg

Museum who kindly re-examined the R ambigua

holotype to confirm my earlier observations. Based

on Hoek’s description, P /aevis can be distinguished

by the more compact trunk and conical proboscis. A

further point of difference is the curvature of the leg

segments. Given that constriction of the scape

segment appears to be a variable character, these

differences are perhaps not significant, but serve to

distinguish the species pending a detailed review of

all available material. Additional figures of P

ambigua are provided.

Pseudopallene watsonae sp. nov.

(Figs 2, 3 J-K)

Pseudopallene ambigua.-Stock, 1973a:

Fig. 7 a, b, d, f. (non P ambigua Stock, 1956),

Material examined

Holotype

SAM E3414, male (ovigerous), Althorpe Is. 35°

22' S 136° 51’ E, Western Isles, S.A., on bryozoan

Orthoscuticella cf ventricosa, 27 m, R. Lewis, 6 Jan

2004.

Allotype

NMV J53155, female, Port Phillip Bay Heads,

Vic., bryozoan Scuticella sp., J. E. Watson, Mar

1980.

Paratypes

NMV J53163, 1 male (ovigerous), Port Phillip Bay

Heads, Vic.,15 — 25 m, J.E. Watson, Mar 1980; NMV

J53164, 1 female, Port Phillip Bay Heads, Vic., 16 m,

D.A. Staples, Nov 1978.

Description

Holotype. Leg span 21.3 mm. Trunk smooth,

segmented, separated from lateral processes by

distinct cuticular line, lateral ecdysial lines well-

defined, arthrodial membrane broad. Neck short,

cephalon with low mid-dorsal mound anterior to

ocular tubercle divided longitudinally by a cuticular

band extending the length of the neck. The band is

darkly pigmented, fading to base of tubercle. Lateral

processes about as long as own maximum diameter,

each with several tiny dorsodistal spines, each

process separated by < '/; own basal diameter,

separation decreasing between posterior processes,

Abdomen slightly inclined, short, inflated, narrows

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 161

Yu)

O.1 mm QS

Fig. 2. Pseudopallene watsonae sp. nov., SAM E3414, male holotype: A, trunk, lateral view; B, trunk, dorsal view; C, trunk,

anterior view; D, third leg; E, tarsus and propodus, third leg; F, oviger; G, oviger claw; H, chela, left , interior; I chela,

left, exterior.

162 D. A. STAPLES

abruptly to a cleft tip, reaching to about half length

of 4th lateral process. Ocular tubercle a little wider

than tall, bearing 2 prominent dorsal papillae, 4

large pigmented eyes of equal size, lateral sense

organs not evident. Proboscis directed ventrally,

mid-region gently curved inward, tapering to oral

surface, conspicuous fringe of dense setae

immediately preceding tip. Scape one-segmented,

little shorter than length of proboscis, slight basal

constriction. Chelae robust, fingers short, tips blunt,

touching when closed; immoveable finger straight,

less that '/s length of palm; moveable finger strongly

curved and rounded, a little off-set distally from

immovable finger, prominent lobe on each finger

corresponds with indentation on inner surface of the

opposing finger. Palm broad, outer surface strongly

inflated, inner and outer surfaces covered in short

setae with a group of longer setae on inner surface

at base of immoveable finger. Oviger segment 5

longest, about 25% longer than segment 4, curved,

distal apophysis prominent, strigilis absent, terminal

claw, distally acute, partially folded over, margins

serrated, distal serrations fine, spine formula

14:10:10:10, proximal spine smallest, distal-most

spine placed dorso-laterally, spines relatively narrow

and long, many partially folded distally, similar to

the main claw, segments 7 — 10 with scattered

surface spinules, bifurcate setae visible under high

magnification.

Measurements

Oviger male (mm). Seg. 1, damaged; Seg. 2, 0.24;

seg. 3, 0.27; seg. 4, 0.95; seg. 5, 1.27; seg. 6, 0.25;

seg. 7, 0.42; seg. 8, 0.26; seg. 9, 0.25; seg. 10, 0.25,

claw 0.17.

Legs slender, scattered short spines on longer

segments, most spines with basal process. Coxa 2 is

2.3 times as long as coxa 3, tibia 2 longest segment,

femur and tibia | subequal, length of tibia 2 about

5.0 times maximum width; tarsus bearing one large

and many smaller spines; propodus weakly curved,

heel low, with 2 (3) large spines of slightly variable

shape followed by two smaller spines in single row,

of the larger spines, the distal-most is largest,

proximal spine slightly curved, sole with median row

of about twelve spines flanked by numerous smaller

spines. Main claw of varying length, from !/2 — */s

length of propodus, Genital pores small and appear

to be on legs 3 and 4 only. Conical area on dorsal

surface coxa 2 of all legs may indicate the presence

of a gland.

Measurements

Holotype (mm). — Length of trunk (frontal margin

of cephalic segment to tip of 4th lateral process),

3.40: width (across 2nd lateral process), 1.50;

proboscis length (lateral), 0.77; Third leg; coxa I,

0.42: coxa 2, 0.97; coxa 3, 0.42; femur, 2.00; tibia 1,

2.0; tibia 2, 2.4; tarsus, 0.22; propodus, 0.75; claw,

0.67.

Allotype

Slightly smaller but otherwise in close agreement

with male. One oviger lost, other, segment 10

damaged, partly missing. Segment 5 marginally

longer than segment 4, segment 4 slightly swollen

proximally. Lateral processes narrowly separated.

Scape with strong proximal constriction. Femora

swollen, each with about 10 eggs. Genital pores

large, all legs.

Variation

The oviger spine formulae are variable, Port Phillip

Bay male, (J53163) 13:10:10:10; female (J53264)

14:9:8:10. The oviger distal claw shape was constant.

Scapes of female (J53164) strongly constricted at

base.

Remarks

L have not examined Stock’s (1973a:115) material

from Pearson I.; however, based on his description

(Figs 7 a, b d, f) it is clear that he has attributed the

juvenile specimens of PR watsonae to P ambigua.

This new material, consisting of mature specimens,

confirms the independent status of R watsonae. The

most conspicuous difference between P watsonae

and P ambigua is in the shape of the chelifore

fingers. Pseudopallene ambigua is further

distinguished by the overall size, being about twice

that of P watsonae and the more slender legs of

which tibia 2 is about 7.2 times as long as the

maximum width. The mid-dorsal region of cephalon

is evenly rounded with no suggestion of a mid-dorsal

process. The proboscis is slightly constricted at about

one-third and two-thirds its length before tapering

distally. The curvature of oviger segment 5 is

stronger and the terminal claw lacks well-defined

teeth. This ovigerous holotype carried several larvae

amongst the eggs. The larvae are hyaline, the 4 legs

each terminated in a claw as long as the propodus

and tarsus together, heel with single spine, propodal

sole with 2 spines (Fig. 3, J). In contrast to the robust

chela fingers of the adult, which are well suited to

crushing bryozoan zooids, the larval chelae are

delicate. The moveable finger is distorted and

digitiform, terminating in a needle-like process

which appears to connect with internal (glandular?)

tissue (Fig. 3 K). The shape of the digitiform process

suggests a specialized purpose, and it is possible that

it may be used to manipulate the manubrium

covering the frontal pore of each zooid facilitating

insertion of the proboscis. Many of the bryozoan

zooids were empty with the operculum of the frontal

pore displaced to a vertical position.

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 163

Fig. 3. Pseudopallene inflata sp.nov., SAM E 3417, female allotype: A, trunk, anterior view; B, trunk, dorsal view; C, trunk,

lateral; D, oviger; E, oviger claw; EF, oviger spines; G, chela; H leg 4; I. tarsus and propodus leg 4; Pseudopallene watsonae

SAM E3414,larval form: J, tarsus and propodus; K, chela.

164 D. A. STAPLES

Etymology: This species is named for Dr. Jeanette

Watson in recognition of her outstanding

contribution to Australian pycnogonid collections.

Pseudopallene inflata sp. nov.

Fig 3

Holotype

SAM E3417, female, Althorpe I., 35° 22’ S; 136°

51' E; 300 m N-NW of The Monuments, on

bryozoan Orthoscuticella cf. ventricosa, 23 m, R.

Lewis, 6 Feb 2004.

Paratypes

SAM E3416, 1 female (subadult), Althorpe Is,

Western Isles, S.A. on bryozoan Orthoscuticella cf

ventricosa, 13 m, D. A. Staples and T. Laperousaz,

6/02/04; SAM E3415, 1 female, eastern Mooring

Bay, Althorpe I. on bryozoan Scuticella, 3-5 m, S.A.

Shepherd, 12/2/05.

Female

(holotype): Leg span 15.5mm. Trunk smooth,

segmented, lateral ecdysial lines well-defined. Neck

short, mid-dorsal mound on cephalon conical,

divided by longitudinal band; dorsal and ventral

surface of all trunk segments inflated, Lateral

processes little longer than maximum width,

separated by about half own basal diameter, each

with several tiny dorso-distal spines. Abdomen short,

rounded, inclined at about 45 degrees. Ocular

tubercle taller than width at base, tapered toward tip,

apical papillae prominent, frontal margin convex; 4

eyes, oval, pigmented; lateral sense organs not

evident. Proboscis directed ventrally slightly

constricted at about one- third and two-thirds length.

Scape strongly constricted proximally, chela fingers

pointed, cutting edges smooth, slightly irregular,

immoveable finger little more than '/2 length of palm,

few setae. Oviger segments 4 and 5 about equal

length, segment 4 swollen proximally, spine formula

segments 7 — 10, 13:9:9:8, terminal claw elongate,

greater than half length segment 10, scoop-shaped,

margins of distal two-thirds bluntly serrated,

serrations larger proximally.

Measurements

Oviger holotype (mm). Seg. 1, 0.10; seg. 2, 0.25;

seg. 3, 0.28; seg. 4, 0.55; seg. 5, 0.58; seg. 6, 0.17;

seg. 7, 0.31; seg. 8, 0.25; seg. 9, 0.23; seg. 10, 0.21,

claw 0.12.

Legs; coxa 2 about 2 times length coxa 3, inflated

distally, tibia 2 longest segment, femur slightly

curved, inflated, surface with scattered spines, little

longer than tibia 1, both tibiae spinous dorsally,

individual spines mounted on small processes,

propodal heel low, 3 - 4 heel spines, distal-most spine

longest, longer than '/2 width of segment, sole armed

with about 10 smaller spines. The remaining third leg

is damaged. Genital pores large, all legs.

Measurements

Holotype (mm) — Length trunk (frontal margin of

cephalic segment to tip of 4th lateral process,

lateral), 1.91; width (across 2nd lateral process), 1.15

proboscis length (lateral), 0.98. 4th leg; coxa I, 0.3;

coxa 2, 0.77; coxa 3, 0.37; femur, 2.09; tibia 1, 2.0;

tibia 2, 2.18 tarsus, 0.12; propodus, 0.67; claw, 0.45.

Variation

The oviger spine formulae variable, distal claw

shapes constant. The spine formulae adult female

SAM E3415 was 12:9:910, SAM 3416 was 11:8:7:7

Remarks

The presence of an acute swelling on the mid

dorsal surface of the cephalon anterior to the ocular

tubercle, together with inflated trunk segments most

readily distinguish this species from its congeners.

The longer heel spines, also, appear to be a useful

diagnostic character.

Etymology

The specific name alludes to the rounded or

inflated trunk segments.

Pseudopallene reflexa (Stock, 1968) comb. nov.

Fig 4, Fig 5 F-G

Spasmopallene reflexa Stock, 1968: 40-42, Fig. 15

a-h.

Type locality

Galathea Stn. 571. Great Australian Bight. (38° 47'

S, 142° 41’ E).

Material examined

SAM E3418, | male (ovigerous), 4 subadults, 3

juveniles; Althorpe Is, Western Isles, S.A. amongst

bryozoan Orthoscuticella cf ventricosa, 13 m, D. A.

Staples and T. Laperousaz, 6 Feb 2004. NMV

J53160, 1 subadult female, Popes Eye, Port Phillip

Bay, Vic., on bryozoan Orthoscuticella ventricosa,

3m, T. O’Hara, 10 Mar 2005.

Description

Male

Leg span 20.0 mm. Trunk smooth, completely

segmented, lateral ecdysial lines not evident. Lateral

processes little longer than maximum width, first

and second pair separated by about two-thirds own

basal diameter, decreasing to about '/s diameter

between segments 3 and 4. Mid-dorsal region of

cephalon, anterior to ocular tubercle, rounded in

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 165

Fig. 4. Pseudopallene reflexa, SAM E3418, male: A, trunk, anterior view; B, trunk, lateral view; C, trunk, dorsal view: D,

third leg; E, tarsus and propodus, third leg; F, oviger; G, oviger claw; H, chela, left interior; I, chela, left exterior.

166 D. A. STAPLES

anterior view, smooth, no evidence of mound or

longitudinal division. Neck short. Abdomen follows

downward curvature of trunk, inflated, barrel-

shaped, cleft at tip, little shorter than distal margin

of 4" Jateral process. Ocular tubercle broader than

tall, four well pigmented eyes, 2 low apical papillae;

lateral sense organs not evident. Proboscis very

slightly inflated at about two-thirds length, almost

parallel sided, narrows sharply distally, dense fringe

of oral setae, proboscis and basal arthrodial

membrane extended. Scape with slight basal

constriction, chelae fingers smooth, pointed,

immoveable finger little longer than moveable

finger and little less than '/2 length of palm; palm

glabrous.

Oviger segment 5 longest, strongly curved, distal

apophysis prominent, acute; segments 4 and 5 with

scattered, fine setae; bifurcate setae visible under

high magnification on segments 4 — 10, spine

formula 12:9:9:8, terminal claw of same form in

adults and subadults, elongate, ladle-like, margin

lined distally with about 20 well-defined teeth,

increasing in length distally.

Oviger measurements

Male holotype (mm). Seg. 1, 0.25; seg. 2, 0.30;

seg. 3, 0.40; seg. 4, 0.90; seg. 5, 1.53; seg. 6, 0.35;

seg. 7, 0.42; seg. 8, 0.30; seg. 9, 0.27; seg. 10, 0.25,

claw 0.18.

Legs; coxa 2, 2.7 times as long as coxa 3, coxa 2

with low, conical swelling on dorsal surface at about

3/4 length of segment, surface of femur and tibiae

with scattered spines, longer spines about '/s segment

width but mostly shorter, dorsal spines particularly

abundant, most surmounted on a basal process,

longer segments uneven, with low swellings on

surface; tibia 2 longest, 4.7 times as long as greatest

width; femur little longer than tibia 1, low swelling

ventrally at '/2 to */3 length, dorsal surface linear,

dorsal surface of tibiae irregular, ventral surface

linear with few spines; tarsus short, numerous

scattered setae on dorsal surface, single larger distal

spine; propodal heel pronounced, spine arrangement

distinctive, typically with two basal median spines,

distal-most spine slightly larger, followed by 4 pair of

slightly smaller spines angled in a ‘V’ arrangement;

sole with about 10 spines. Main claw little less than

2/3 length of propodus. Genital pores small, ventral

surface coxa 2 legs 3 and 4. Auxiliary claws absent.

Measurements

(male mm). Length trunk (frontal margin of

cephalic segment to tip of 4th lateral process), 3.55;

width (across 2nd lateral process), 1.42; proboscis

length (lateral), 1.37. Third leg; coxa 1, 0.37; coxa 2,

1.32; coxa 3, 0.50; femur, 2.00; tibia 1, 1.70; tibia 2,

2.37; tarsus, 0.22; propodus, 0.65; claw, 0.47.

Remarks

These specimens have taken up the orange-brown

colour of the gut contents and match the Scuticellid

bryozoan on which they were found. The juvenile

specimen is in close agreement with the form

described as Spasmopallene reflexa Stock, 1968 first

described from the region. In particular, it agrees in

the shape of the proboscis and distal oviger claw.

Unfortunately, however, | have been unsuccessful in

confirming the characteristic arrangement of heel

spines on the holotype. Stock recorded 4 — 5 heel

spines but it is possible that 4 of the spines observed

by Stock obscured matching pairs and that the 5th

spine represents a median basal spine. Based on

Stock’s description, and the material before me, I

have little reason to doubt that the same species is

represented. The 2nd coxae of subadults and

juveniles are short, 1.6 — 1.8 times longer that coxa

3, and the propodus is much more strongly curved

with a more prominent heel. The gut follows the

contours and irregular surface of the legs, narrowing

and dilating intermittently giving it a globular

appearance. The proboscis of one subadult female

tapers from about !/2 its length, distal portion plug-

like, expanding slightly before again narrowing to

setose tip (Fig. 5 G). Pseudopallene reflexa is readily

distinguished from other described species by the

irregular surface of the longer leg segments, the

distinctive paired arrangement of the propodal heel

spines, the proboscis shape, the stronger curvature of

the 5th oviger segment and the form of the terminal

oviger claw.

Distribution

Bicheno, Tas.; Bass Strait; Port Phillip Bay, Vic. at

depths 3 — 72 m.

Stylopallene Clark, 1963

Stylopallene cheilorhynchus Clark, 1963

(Fig 5 A, B)

Stvlopallene cheilorhynchus Clark, 1963: 36-38,

Fig. 19 A-I.? Stock, 1973a: 117.?Stock, 1973b: 92.

Type locality

Port Arthur, Tasmania.

Material examined

SAM E3420, 2 subadults, 1 larval form, Althorpe

Is, 300 m N-NW of The Monuments, S.A. 23 m, R.

Lewis, 6 Feb 2004; SAM E3421, 5 immature

specimens, 2 larval forms, Western Isles, 27 m, R.

Lewis, 6 Feb 2004; SAM E3422, 7 males, 4 females,

6 subadults, | larval form, Western Isles, on

bryozoan Amathia wilsoni, 13 m, D. Staples and T.

Laperousaz, 6 Feb 2004; SAM E3423, 1 male,

eastern Mooring Bay, Althorpe L, 3 — 5 m, S.A.

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 167

Fig. 5. Stylopallene cheilorhynchus SAM E3422: A., trunk, male, lateral: B, chela, juvenile. Callipallene micracantha SAM

£3424, male, C, oviger spines; D, leg 3; SAM E3425, female, E, leg 3; P. reflexa, SAM E3418, F, propodus, juvenile leg

3; G, proboscis, sub adult.

Shepherd, | 2/02/05; SAM E3429, | male ovigerous,

Nora Creina, S.A., snorkel collection, A.I.M.S

collecting Team, 20 Feb 1989; SAM E3438, 1

subadult, Cape Northumberland, S.A. in algae, 15 m,

S.A. Shepherd, Feb 1977; SAM E3430, | male

ovigerous, St Francis I., off SW point, S.A., rocks

and sand 35 m, W. Zeidler, 29/01/1982; SAM E3431,

1 female, Pearson I., S.A., 30 — 33 m, (no additional

data); SAM E3432, 9 larval forms and juveniles, off

Cape Jaffa, approx 3 km WNW of Margaret Brock

Reef Lighthouse, S.A., ridges crevices, overhangs,

algae, few sand pockets, W. Zeidler and K.L.

Gowlett-Holmes, 17 Feb 1989; SAM E3433, 2

females, Dutton Cove, NE Kangaroo [., S.A. in night

time plankton tow, J. Ottaway, 19 Jan 1979; SAM

E3428, | male ovigerous, Cape Thomas, between

Godfrey I., S.A., 3 — 7 m, red algae, soft rock, reef,

sand wreck, W. Zeidler and K. Gowlett-Holmes, 16

Feb 1989; SAM E3434, 1 female, Kingscote,

Kangaroo I., S.A., 5 m, on hydroids, N. Coleman, 13

Mar 1978; SAM E3435, | Juv., Robe, S.A., | km off-

shore, 15 m, D. Staples, 16 Apr 1976; SAM E3436,

| male ovigerous, West I., Encounter Bay, in

sortings, 12 m, J. Ottaway, 26 Nov 1978; SAM

E3437. 1 male ovigerous, Hopkins |., SE Eyre

Peninsula, S.A., 17 m, 30 Nov 1995.

Remarks

A compact species, easily confused with closely

related but more elongate, S. longicauda Stock,

1973a, and shares similar body markings. The distal

'o of the longer segments of the legs is black,

contrasting against the proximal off-white colouring

giving a banded appearance to the specimens. There

168 D. A. STAPLES

is little to add to Clark’s (1963) description. Clark

described the femur and tibia 2 as subequal; in this

material tibia 2 is consistently about 1.2 times longer

than the femur. Female genital pores are prominent

and swollen. Male genital pores are tiny and obscure

in comparison. Additional figures showing the adult

in lateral view and juvenile chela are provided. This

species was present in vast numbers on colonies of

the arborescent bryozoan Amathia wilsoni. The large

proportion of egg-bearing adults and the presence of

so many immature forms may indicate a summer

breeding season.

Distribution

St Francis Isles, S.A. to Coffs Harbour, N.S.W. and

Tasmania. Low-water to 90 m. Very common in SE

Aust.

Callipallene Flynn, 1929

Callipallene micracantha Stock, 1954

Fig 5 C-E

Callipallene emaciata micracantha Stock, 1954:

44-46, figs 19, 20 a-b, — Clark, 1963: 21-23, Fig. 11

A-I. — Child, 1975: 10.

Type locality

Off Cape Everard, Victoria (37° 05’ S, 150° 05’ E).

Material examined

SAM _ E3424, 1 ovigerous male, Althorpe Is.,

Western Isles, S.A., 27 m, R. Lewis, 6 Feb 2004;

SAM E3425, 2 males, | female, Backstairs Passage,

J. E. Watson, (undated); SAM E3439, 1 female, Cape

Northumberland, S.A. in algae, 15 m, S.A. Shepherd,

1 March 1975; SAM E3440, | female, Gulf St

Vincent, S.A. on Posidonia sinuosa, S.A. Shepherd,

May 1985; SAM E3441, 1 female, Wallaroo,

Spencer Gulf, S.A., J. E. Watson, Jan 1983.

Remarks

These specimens substantially agree with Stock’s

(1954) description of the male holotype. Stock

erected the subspecies micracantha to accommodate

a single male specimen from eastern Bass Strait,

which differed from other species and subspecies of

the genus in the spination of the lateral processes and

crop. His specimen was also notable in the presence

of a double row of denticles on the immoveable

finger, a character shared with C. cuspidata Stock,

1954 and Neopallene antipoda Stock, 1954. The

double row of denticles is not present in this male

specimen; however examination of additional

material from the Victorian coastline and Bass Strait

indicates this to be a variable character in adults of

both sexes. The ventral spine-tipped swellings of the

femur and to a lesser extent tibia 1, are present in all

southern Australian specimens but more pronounced

in the female, particularly egg-bearing specimens

(Fig. 5, E). The crenulate propodal spines and

serrated auxiliary claws characteristic of C. emaciata

(Child 1979:41) are not present in the material

examined. In the shape of the femur and tibia | the

Australian material agrees with C. emaciata subspec.

(Stock, 1954:46) from New Zealand but significant

differences in the length of the neck and shape of the

proboscis distinguish the species. Stock (1954) was

reluctant to give his specimen species status on the

basis of his solitary male, but given the wide range of

material now available and the consistency in

significant morphological characters, specific rank is

well justified. The oviger spines are dimorphic. The

distal-most spine on oviger segments 7 — 9 is off-set

and larger than preceding spines. The teeth are

irregular, well defined and larger on the distal

margin. Dimorphism is not unique in the genus but

in general shape, the distal spine morphology is close

to that found in C. panamensis Child, 1979, C.

brevirostris novaezealandiae (Thomson, 1884) and

C. phantoma (Dohrn, 1881). Spines preceding the

distal spine are uniform and finely denticulate.

Spines at the base of the chelifore implants are

variable in number, 3-4 most common, often with

about 3 additional spines on the lateral margin.

Female genital pores are larger than those of the male

and present on all legs.

Distribution

Pearson I., S.A. to Batemans Bay, NSW and

Tasmania 10 — 135 m

AMMOTHEIDAE Dohrn, 1881

Achelia Hodge, 1864

Achelia transfugoides Stock, 1973

Achelia transfugoides Stock, 1973a: 104-106, Fig.

2. — Stock, 1973b: 92.

Type locality

Toad Head, West I., South Australia.

Material examined

SAM E3426, 1 female, Althorpe Is, Western Isles,

S.A., in sortings, 13 m, D. Staples and T. Laperousaz,

6 Feb 2004: SAM E3427, | male, eastern Mooring

Bay, Althorpe I., S.A. on bryozoan Amathia sp, 3 —5

m, S.A. Shepherd, 12/02/05; SAM E3442, 1

subadult, 7 females, 5 males ovigerous, Fanny Point,

Boston I., Spencer Gulf, S.A. on Sargassum, reef, 2-

8 m, W. Zeidler, K.L. Gowlett-Holmes, 17 Feb 1988,

Remarks

| have tentatively assigned this material to A.

transfugoides, the principal difference being in the

PYCNOGONIDA FROM THE ALTHORPE ISLANDS, SOUTH AUSTRALIA 169

shape of the proboscis. Stock (1973a:104) described

the proboscis with a swollen basal part which is

consistent with his Fig. 2 c showing maximum width

at the base. This figure is however, inconsistent with

his figures b, e, in which he shows the basal part

narrower than the inflated mid- region and with

which the present material agrees. The tubiform part

of the proboscis is a little shorter than illustrated by

Stock (Fig. e), and in some other material I have

examined, but otherwise closely agrees with the type

material.

Distribution

Perth, WA to Wilsons Promontory, Vic.

Acknowledgements

I am grateful to the scientific party and crew of the

RV Ngerin for their assistance and support. In

particular, | thank Sue Murray-Jones for facilitating

my participation, and fellow divers and collectors,

Thierry Laperousaz, Scoresby Shepherd, Rob Lewis,

and James Brook. Special thanks to H. Dastych of the

Zoological Museum Hamburg for lending the

holotype material, to Robin Wilson and Angelika

Brandt for transporting the specimens, and to Phil

Bock for identifying bryozoans. Finally I acknowledge

the helpful criticism of G.C.B.Poore, and of the two

reviewers, Franz Krapp and Roger Bamber.

References

Cuitp, C. A. (1975) Pycnogonida of Western Australia.

Smithson. Contr. Zool. 19, 1-29.

(1979) Shallow-water Pycnogonida of the

Isthmus of Panama and the coasts of Middle America.

Smithson. Contr. Zool. 293, 1-86.

Crark, W. C. (1963) Australian Pycnogonida. Rec. Austr:

Mus. 26 (1), 1-81.

Donen, A. (1881) Die Pantopoden des Golfes von Neapel

und der angrenzenden Meeresabschnitte. Fauna Flora

Golfe. Neapel 3, 1-252.

Hoek, P. C. C. (1881) Report on the Pycnogonida dredged

by HMS Challenger 1873-76. Rep. Scien Results Explor

Voyage Challenger 3 (10), 1-167.

Murray-JONnes, S. & SHEPHERD, S. A. (2005) Introductory

narrative and conservation recommendations. Trans. R.

Soc. S. Aust. 129, 85-89.

SCHIMKEWITSCH, W. (1909) Nochmals tiber die Periodizitit

in dem System der Pantopoden. Zool. Anz. 34 (1), 1-13.

StapLes, D. A. (1997) Pycnogonids. pp.1040-1072 In

Shepherd, S. A. & Davies, M. (Eds) “Marine Invertebrates

of Southern Australia Part 111’. (SARDI and Fauna and

Flora of SA Handbooks Committee, Adelaide).

Stock, J. H. (1954) Pycnogonida from Indo-West Pacific,

Australian and New Zealand waters. Vidensk Meddr

dansk naturh. Foren. 116, 1-168.

(1956) Pantopoden aus dem Zoologischen

Museum Hamburg. Mitt. Hamb. zool. Mus. Inst. 54, 33-

48.

(1968) Pycnogonida collected by the

Galathea and Anton Bruun in the Indian and Pacific

Oceans. Vidensk. Meddr dansk naturh. Foren. (131),

7-65.

(1973a) Pycnogonida from south-eastern

Australia. Beaufortia 20, (266), 99-127.

(1973b) Achelia shepherdi n sp and other

Pycnogonida from Australia. Beaufortia. 21, (279), 91-

THomson, G. M. (1884) On the New Zealand Pycnogonida,

with descriptions of new species. Trans. Proc N. Z. Inst.

16, 242-248.

WILson, E. B. (1878) Descriptions of two new genera of

Pycnogonida. Am. J. Sci. 15, 200-203.

Transactions of the Royal Society of S. Aust. (2005), 129(2), 170-182.

A NEW SPECIES OF NEOPELTOPSIS (COPEPODA, HARPACTICOIDA,

PELTIDITIDAE) FROM ALTHORPE ISLAND, SOUTH AUSTRALIA

by G. K. WALKER-SMITH!

Summary

WALKER-SMITH, G. K. (2005) A new species of Neopeltopsis (Copepoda, Harpacticoida, Peltidiidae) from

Althorpe Island, South Australia. Trans. R. Soc. S. Aust. 129(2), 170-182, 30 November, 2005.

A new species of Neopeltopsis Hicks, 1976 is described from algae collected from Althorpe Island, South

Australia. The new species is separated from its congeners by a number of morphological characters related to

the A2, mandible, P1, P2, PS and the male Al. A revised generic diagnosis for Neopel/topsis is provided and

intra- and intergeneric relationships are discussed. The occurrence of other species of Peltidiidae, collected from

algal washings from Althorpe I., is documented. Neopeltopsis appears to be restricted to the Southern

Hemisphere, with previous records limited to New Zealand (N. pectinipes Hicks, 1976) and Argentina (N. hicksi

Pallares, 1979).

Key Worns: Algae, Harpacticoida, Neopeltopsis, new species, Peltidiidae, South Australia.

Introduction

Harpacticoid copepods are microcrustaceans,

usually less than 1 mm in length, that occur in

marine, estuarine, and freshwater habitats. There are

approximately 50 families and 460 genera of

Harpacticoida and estimates of the total number of

described species range between 3000 (Huys et al.

1996) and 4000 to 4500 (Giere 1993).

The Australian harpacticoid fauna is abundant and

diverse, but remains largely undescribed. One

hundred and thirty-three species of Harpacticoida

have been recorded in Australia, with major

contributions being made by Nicholls (1941, 1942,

1945a-c, 1957), Hamond (1971, 1973a-e, 1974,

1987), Harris (1994, 2002), Harris & Robertson

(1994), Bartsch (1993, 1994, 1995, 1999) and

Karanovic (2004). Thirty-one species have been

recorded from South Australia, including 11

Australian endemics described by Nicholls (1941,

1942, 1945a) and Hamond (1971, 1973c, 1973e).

In February 2004, algal samples were gathered

from Althorpe L, South Australia (SA) and

harpacticoids living among the thalli were extracted.

Among these harpacticoids, a new species of

Peltidiidae Sars, 1904 belonging to the genus

Neopeltopsis Hicks, 1976 was _ discovered.

Worldwide, the family Peltidiidae consists of eight

genera and 58 species. Species in this family are

typically algal-dwellers, with dorsoventrally

flattened bodies and modified appendages that

' Marine Invertebrate Section, South Australian Museum, North

Terrace, Adelaide, South Australia 5000, Australia, Present

address: School of Zoology, University of Tasmania, Private

Bag 5, Hobart Tasmania 7001 and The Tasmanian Museum

and Art Gallery, GPO Box 1164, Hobart Tasmania 7001.

Email: geneforw@postoffice.utas.edu.au

enable them to exist on the surface of the algal thalli

(Hicks 1986). Prior to the present study, six species

of Peltidiidae were known to occur in SA: Alteutha

depressa (Baird, 1837); Alteutha spinicauda

Nicholls, 1941; Parapeltidium cristatum Nicholls,

1941; Parapeltidium dubium Nicholls, 1941;

Peltidium proximum Nicholls, 1941 and Peltidium

simplex Nicholls, 1941. Peltidium speciosum

Thompson & A. Scott, 1903, was recorded in South

Australia by Nicholls (1941); however, the validity of

this species is doubtful; Wells & Rao (1987) believed

it could not be distinguished from several other

species of Peltidium Philippi, 1839. The only other

species of Peltidiidae recorded from Australia is

Alteuthellopsis corallina Humes, 1981, which was

found associated with scleractinian corals in

Queensland (Humes 1991).

This paper provides the description of a new

species of Neopeltopsis, collected from Althorpe I.,

as well as a revised diagnosis of the genus.

Neopeltopsis currently comprises only two species:

N. pectinipes Hicks, 1976, recorded from New

Zealand and N. hicksi Pallares, 1979 from waters off

Argentina. The new species, described here, is

separated from its congeners by morphological

characters related to the A2, mandible, Pl, P2, P5

and the male Al. This paper also documents the

occurrence of other species of Peltidiidae found in

algae collected from Althorpe I., SA.

Material and Methods

Algae were collected by hand. Algal samples were

washed in a bucket of freshwater and this water was

then poured through a 63 4m mesh sieve. Retained

material was fixed in 95% ethanol and _ later

transferred to 70% ethanol. Samples were examined

NEW SPECIES OF NEOPELTOPSIS 171

under a Wild MB8__ stereomicroscope and

harpacticoids were extracted using fine forceps.

Harpacticoids were dissected in a drop of glycerol on

a microslide using electrolytically-sharpened

tungsten needles. Appendages were transferred to

new microslides, mounted in Gurr’s Aquamount and

coverslips were sealed with clear nail varnish.

Microslides were examined using either a Leitz

Dialux 22 compound microscope with phase contrast

or a Leica DMR compound microscope with

interference contrast. Illustrations were made with

the aid of a camera lucida. Selected specimens were

examined using a Philips XL20 scanning electron

microscope (SEM). These specimens were

dehydrated in a graded ethanol series, critical point

dried using CO, and gold coated prior to examination

under the SEM (K V=10, spot size 3).

Terminology used follows Huys & Boxshall

(1991). Abbreviations used are: Al, antennules or

first antennae; ae, aesthetasc; A2, antennae or second

antennae; Md, mandible; Mxl, maxillule; Mx,

maxilla; Mxp, maxilliped; P1 — P4, swimming legs

1 — 4. Individual segments of Pl — P4 rami are

written (for example) as Pl exopod-3, for the third

(or terminal) segment of the Pl exopod. P5 and P6

refers to the fifth and sixth leg respectively. Total

length measurements are from the tip of the rostrum

to the posterior margin of the caudal rami (excluding

caudal setae). Armature formulae for swimming legs

are constructed following Lang’s (1934) method

(also see Huys & Boxshall 1991: 29), The term

“armature” refers collectively to articulating

elements such as setae and spines. All material

examined is held in the collections of the South

Australian Museum (SAM) and the Tasmanian

Museum and Art Gallery (TMAG).

Neopeltopsis Hicks, 1976

Neopeltopsis Hicks 1976: 363-370.—Hicks 1986:

356, 360-361.

Diagnosis

Body distinctive, broad, dorsoventrally flattened

with simple pattern of chitinous thickening. P1-

bearing somite incorporated into large cephalosome;

P2 — PS somites free, epimeral plates well-

developed; remaining abdominal somites fused,

much shorter than prosome. Anal somite and caudal

rami free. Urosome-caudal rami complex analogous

to that of Porcellidium (Porcellidiidae Sars, 1904)

(Hicks 1986). Rostrum broad, prominent, not

defined at base; 2 Al 8-segmented, d Al 8 — 9

segmented. A2 exopod reduced, 1- or 2-segmented,

with 2 or 4 setae in total; basis without, and endopod-

1 with, abexopodal seta; endopod-2 with 1 lateral

and | distal large, pectinate (comb-like) seta.

Mandibular gnathobase elongate, palp with 1-

segmented exopod and endopod. Maxillule arthrite

(i.e. praecoxal endite) with 6 — 8 spinose spines on

distal margin and 2 setae on anterior surface; coxal

endite with 2 — 3 setae; basis elongate; exopod

distinct, 1-segmented, with 2 — 3 setae; endopod

incorporated into basis and represented by 2 — 3

setae. Maxilla with 3 syncoxal endites, proximal

most widely separated, with 4, 2 and 3 setae

respectively; endopod fused to basis (forming

allobasis), endopod represented by 2 setae, allobasis

with | spinose and 2 naked setae distally. Maxilliped

subchelate; pedestal well-developed; syncoxa armed

with | — 2 setae, basis expanded, ovoid with pad-like

distal seta; endopod drawn out into elongate, narrow

claw, sometimes with small setae on lateral surface

of claw. Pl coxa and basis elongate, orientated at

right angles; exopod 3-segmented, exopod-3

reduced, indistinctly separated from exopod-2,

bearing 4 broadly flattened, pectinate setae, without

accessory armature, geniculate seta absent; endopod

2-segmented, reduced, much shorter than exopod. P2

— P4 rami 3-segmented. Armature formulae for both

sexes as follows:

Exopod Endopod

p2 0.1.22 (1 —2) 0—1.0— 1.1 — 2) 20

P3 0.1.322 1.1.220

P4 0.1.322 1.1.220

P5 both sexes 2-segmented or sometimes

indistinctly 2-segmented. Genital double somite and

distal somites fused, expanded posterolaterally and

almost surrounding the caudal rami. 2 genital

apparatus comprising paired genital apertures

located ventrally on urosome, apertures covered by

P6; copulatory pore located on ventral midline

(slightly) posterior to genital apertures, covered by

operculum. Eggs in single egg-sac. Caudal rami

short and subrectangular with 7 setae, principal

terminal setae distinct (i.e. not fused to one another

at base). 5 P6 left and right identical, large, lobe

shaped; not known for N. pectinipes or N. hicksi.

Male with 1(?)—2 spermatophores (not known for N.

hicksi).

Sexual dimorphism in body size, Al, PI and PS.

Species

Neopeltopsis pectinipes Hicks, 1976; N. hicksi

Pallares, 1979; Neopeltopsis sp. nov. described

herein.

Distribution

Wellington, New Zealand (Hicks 1976); Argentina

(Atlantic Ocean and Tierra del Fuego) (Pallares

1979); Althorpe I., SA.

172 G. K. WALKER-SMITH

Fig. 1. Neopeltopsis althorpensis sp. nov.: A, 2 habitus, dorsal (holotype, SAM C6219). B, d habitus, dorsal (paratype,

SAM C6220). Terminal caudal setae cut short. CO = copulatory operculum, CP = copulatory pore, SR = seminal

receptacle, P6 = sixth leg, S = spermatophore.

Habitat

Phytal. Recorded from: Pterocladia lucida,

P. pinnata and Caulerpa brownii in New Zealand;

Macrocystis (Phaeophyta) and species in the

family Delesseriaceae (Rhodophyta) off the coast

of Argentina; Pterocladia sp. and Lobospira

bicuspidata in SA.

Remarks

Herein the generic diagnosis of Hicks (1986) has

been expanded to better define the genus.

Neopeltopsis althorpensis sp. nov.

(Figs 1 — 8)

Material examined

Holotype. Althorpe I., SA (35° 22.02’ S, 136°

51.08’ E), from washings of Pterocladia sp.

(Rhodophyta), depth ~2 m, coll. A. J. Hirst, 01 Feb.

2004, SAM C6219 (ovigerous & , dissected, mounted

on 9 slides).

Paratypes

Collected with holotype. SAM C6220 (1 6 in

ethanol, allotype); SAM C6221 (1 6, dissected,

mounted on 3 slides. P3 and P4 lost); TMAG G5474

(1 6, partially dissected, in ethanol; P2 mounted on

1 slide); TMAG G5475 (1 2, mounted on | slide);

SAM (C6222 (2 ovigerous 2°, 4 dd, | juv., in

ethanol); TMAG G5476 (2 ovigerous ° 2, | non-

ovigerous 2, 3 od, 5 juv., in ethanol).

Other material

Collected with holotype. SAM C6223 (1 2

mounted on SEM stub), SAM C6224 (2 36,

mounted on SEM stub).

NEW SPECIES OF NEOPELTOPSIS 173

a a

“<<

Coo

Fig. 2. Neopeltopsis althorpensis sp. nov.: A, 2 A1, dorsal (holotype, SAM C6219). B, ¢ Al, dorsal (paratype, SAM

C6220); C, 3 Al, ventral (paratype, SAM C6220).

174 G. K. WALKER-SMITH

Ss EF

D {0.05 mm

0.05 mm

Fig. 3. Neopeltopsis althorpensis sp. nov., 2° holotype, SAM C6219: A, A2; B, Md; C, Mxl; D, Mx; E, Mxp, anterior; F,

Mxp, posterior (basis and endopod only).

NEW SPECIES OF NEOPELTOPSIS 175

Diagnosis

Al & with curved, thorn-like projections on

segment 7. A2 exopod |-segmented, with 4 setae.

Mandible exopod length approximately equal to

width. Pl exopod-2 0.25 size of exopod-l. P2

endopod-2 without inner seta. P2 exopod-3 with only

1 outer spine. P5 exopod partially fused to

baseoendopod in both sexes.

Description of female

Mean total body length 0.86 mm + 0.09 mm

(n = 3). Body dorso-ventrally flattened, simple

pattern of chitinous thickening (Fig. 1A), integument

with numerous pores and sensillae (as found in male,

see Fig. 7B), Caudal rami short and subrectangular,

with 7 setae. Rostrum broad and prominent, fused at

base (Fig. 1A). Caudal rami (Fig. 6C) length ~

width; inner distal margin finely serrate; with 7

setae, seta I shortest, seta IT dorsal to seta I, seta V

well-developed and pinnate, seta VII dorsal and

triarticulate at base.

Al 8-segmented (Fig. 2A); all seta bare. Segment

| with | seta and few fine setules; segment 2 with 11

setae and some short setules; segment 4 with

aesthetasc fused basally to 1 seta; segment 8 with a

shorter aesthetasc fused basally to 2 setae (i.e. a

acrothek), Armature formula for Al: 1-[1], 2-[11], 3-

[8], 4-[3+ (I+ae)], 5-[2], 6-[2], 7-[2], 8-[3+acrothek]

A2 (Fig. 3A) basis and endopod-1 separate; basis

with few fine setules and unarmed; endopod-1 with

| abexopodal seta; endopod-2 with 2 surface frills

distally; lateral armature consisting of | pectinate

spine, | spine and 2 setae basally fused; distally with

I pectinate spine and 4 geniculate setae (innermost

basally fused to naked seta); exopod 1-segmented,

with 4 bare setae.

Labrum and paragnaths not illustrated.

Mandible (Fig. 3B) coxa narrow and elongate,

expanding distally to small gnathobase; basis with |

seta; exopod |-segmented, small (about as long as

wide), with 3 terminal setae; endopod with 1 short

and 3 long setae distally and 1 proximal seta. All

setae naked.

Maxillule (Fig. 3C) arthrite of praecoxa with 8

pinnate and 2 smooth setae; no setae observed on

medial surface; coxal endite with 2 bare setae; basis

elongate, with 5 smooth setae, | geniculate seta and

I serrate spine; exopod |-segmented, with 2 smooth

setae; endopod completely incorporated into basal

segment, but represented by 2 setae.

Maxilla (Fig. 3D) syncoxa with 3 endites; proximal

endite with 4 setae; middle and distal endites with 2

spinulose setae and 3 bare setae respectively;

allobasis bearing | pinnate spine and 2 naked setae;

endopod completely incorporated and represented by

2 setae.

Maxilliped (Figs 3E — F, 8A) subchelate; pedestal

well-developed; syncoxa smallest segment bearing 2

plumose setae; basis ovoid with a short, stout, pad-

like seta with tiny spinules (arrowed in Fig. 8A); with

few spinules proximally along the palmar margin;

endopod drawn out into recurved claw, as long as

basis, 2 small setae on lateral surface of the claw.

Pl (Figs 4A, 8C). Coxa without armature. Basis,

inner seta inserted above endopod; with few spinules

around insertion of outer basal seta and | tube pore

on outer anterior surface and patch of tiny spinules as

figured. Exopod 3-segmented; exopod-3 small and

indistinctly separated from exopod-2; exopod-1 with

smooth outer seta and patch of tiny denticles as

figured; exopod-2 with bare outer seta dorsolaterally

and inner seta strongly serrate and posteriorly

displaced, with a patch of tiny denticles on the dorsal

surface (arrowed in Fig. 8C); exopod-3 with 4

strongly pectinate flattened setae. Endopod reduced,

2-segmented; endopod-1 ~ 4 x length of endopod-2,

without setae; endopod-2 tiny, with 3 bare setae.

P2 — P4 (Figs 4B, SA — B) rami 3-segmented.

Armature formulae as follows:

Exopod Endopod

P2 0.1.221 1.0.220

P3 0.1.322 1.1.220

P4 0.1.322 1.1.220

P2 (Fig. 4B). Praecoxa and coxa not illustrated,

coxa with anterior pore. Basis elongate with bare

outer seta. Exopod-1! with fine setae on inner margin

and spinulose spine on laterodistal corner; exopod-2

with plumose inner seta and pinnate outer spine;

exopod-3 shorter than 2 preceding segments, with 2

inner, plumose setae, 2 distal setae with spinules and

setules and | pinnate outer spine. Endopod-1 with |

inner seta and fine setules along outer margin;

endopod-2 without setae; endopod-3 with 2 inner

setae and 2 distal setae.

P3 (Fig. SA). Praecoxa and coxa not illustrated.

Basis transversely elongate, with bare outer seta.

Exopod-1 with patch of fine spinules around distal

outer corner, spinulose spine, no inner seta; exopod-

2 with plumose inner and pinnate outer seta, with |

anterior and 2 posterior spinule rows and a pore in

the distal margin of the segment (arrowed in Fig.

5A); exopod-3 with 3 inner plumose setae (2

proximal most with additional pinnules), 2 distal

setae with pinnules and setules, 2 pinnate outer

spines, 2 spinule rows posteriorly. Endopod-1 with |

plumose inner seta; endopod-2 with inner seta

(broken off in holotype, present in paratype, position

stippled in Fig. 5A); endopod-3 with 2 inner setae

and 2 distal setae.

P4 (Fig. 5B). Praecoxa and coxa not illustrated.

G. K. WALKER-SMITH

176

pee LILES se

FP LAP

Fig. 4. Neopeltopsis althorpensis sp. nov., 2 holotype, SAM C6219: A, PI anterior; B, P2 anterior.

177

S OF NEOPELTOPSIS

NEW SPECIE

Fig. 5. Neopeltopsis althorpensis sp. nov., 2 holotype, SAM C6219: A, P3 anterior; B, P4 posterior. Arrows indicate pore.

178 G. K. WALKER-SMITH

Chee SS St

Fig. 6. Neopeltopsis althorpensis sp. nov.: A, 2 P5 anterior (holotype, SAM C6219); B, ¢ PS anterior (paratype, SAM

C6221); C, 2 right caudal ramus, dorsal (holotype, SAM C6219), setae numbered; D, ¢ right caudal ramus, dorsal

(paratype, SAM C6220).

NEW SPECIES OF NEOPELTOPSIS 179

Basis elongate with | naked outer seta. Exopod-1

with pinnate outer spine; no inner seta; exopod-2

with 2 rows of small spinules, outer spine pinnate

and elongate, nearly twice as long as entire exopod-

2, inner seta plumose, 2 rows of spinules on posterior

surface and a pore in the distal margin of the segment

(arrowed in Fig. 5B); exopod-3 with 3 inner setae

(proximal with additional row of pinnules), 2 distal

setae, 2 pinnate outer spines, 2 inner, posterior

spinule rows. Endopod-! with | plumose inner seta;

endopod-2 without seta; endopod-3 with 2 inner

setae and 2 distal pinnate setae.

P5 (Fig. 6A) exopod partially fused to

baseoendopod. Baseoendopod outer seta smooth and

arising from setophore; endopodal lobe with 4 bare

setae and | tube pore. Exopod with 5 pinnate setae,

1 naked seta and | ventral pore.

Po (Fig. 1A) small, kidney shaped, with 1 seta;

covering genital apertures. Single copulatory pore

(Fig. 1A) located on ventral midline (slightly)

posterior to genital apertures, covered by operculum

(Fig. 1A).

Description of male

Mean total body length 0.68 mm + 0.05 mm

(n= 5). Body (Figs 1B, 7A). Sexual dimorphism in

body size, Al, Pl and P5. Al (Figs 2B — C, 7C)

haplocer, 8-segmented, segment 7 with 2 curved,

thorn-like projections, aesthetascs on segments 3

and 5. Armature formula for Al: 1-[1], 2-[11],

3-[5 + (1+ae)], 4-[2], 5-[5 + (1+ae)], 6-[0], 7-[0],

8-[9]. Oral appendages as in female. Swimming

legs as for female except: dP1 proportionally

longer than that of the female; extends to distal

edge of caudal rami whereas 2 PI only reaches to

distal margin of egg sac. P5 (Fig. 6B) exopod

partially fused to baseoendopod; outer basal

setophore bearing | naked seta; endopodal lobe

with | naked seta and | tube pore; exopod with 3

smooth and 3 spinulose setae; ventral exopodal

surface with pore. P6 (Fig. 1B) left and right

identical, large, semi-circular, without setae.

Urosome with 2 spermatophores.

Etymology

The specific name althorpensis is derived from the

type locality, Althorpe L, SA.

Variability

The left P2 endopod-2 and endopod-3 of the

holotype were partially fused along the inner margin

but the right P2 endopod was clearly 3-segmented,

which is the normal condition. The left PS of one of

the male paratypes was smaller than the right one;

the right PS represented the normal size as it was the

same size as both the left and right P5 in other male

paratypes.

Remarks

This species was found (rarely) on Lobospira

bicuspidata (Phaeophyta).

The family level diagnosis of Huys et al. (1996)

and Boxshall & Halsey (2004) must be emended as

N. althorpensis and N. pectinipes males have two

spermatophores, not one.

Discussion

Neopeltopsis is distinguished from other

Peltidiidae genera by two autapomorphies: the

possession of four pectinate setae on the Pl exopod-

3 and the medial fusion of the abdominal somites. No

other characters are unique to just one genus of

Peltidiidae (Hicks 1986), and for this reason

Neopeltopsis can be considered the best defined

genus of the Peltidiidae. The possession of five

armature elements on the Pl exopod-3 is considered

the plesiomorphic condition and while there are

other genera of Peltidiidae with four armature

elements on the terminal segment of P1, they are not

pectinate (i.e. Peltidium and Parapeltidium A. Scott,

1909). As the new species from Althorpe I. possesses

four pectinate setae on the P] exopod-3 and exhibits

fusion of the abdominal somites, it is placed in the

genus Neopeltopsis. In addition to these two

apomorphies, Neopeltopsis shares many other

character states with N. hicksi and N. pectinipes;

however, it is separated from its congeners by:

1) Al of male with two curved, thorn-like

projections on segment 7 (N. pectinipes with one

thorn-like projection on segments 6 and 7; N.

hicksi has only one projection on segment 7).

Based on the illustrations of Hicks (1976) and

Pallares (1979) it is assumed that the

segmentation of the Al of the male was

misinterpreted. It is most probable that segment

4, a small sclerite, was mistakenly considered to

be part of segment 3. It is also possible the

terminal segment observed by Hicks (1976) is in

fact fused to the preceding segment. If this is not

the case, and Hicks (1976) did overlook the small

somite which is segment 4, then the dAI of

N. pectinipes has 10 segments;

2) A2 exopod l-segmented and with 4 setae (N.

pectinipes \-segmented with 2 setae; N. hicksi

2-segmented, endopod-! with | seta and

endopod-2 with 3 setae);

3) Mandible exopod length ~ width (N. pectinipes

length ~2 x width, i.e. cylindrical; NV. hicksi same

as for N. althorpensis);

4) Pl exopod-2 0.25 size of exopod-1 (N. pectinipes

0.75 size of exopod-1; N. hicksi approximately

equal to exopod-1);

5) P2 endopod-2 without inner seta (N. pectinipes

and WN. hicksi with | inner seta):

180 G. K. WALKER-SMITH

Fig. 7. Neopeltopsis althorpensis sp. nov., 5 paratype, SAM C6224: A, habitus, dorsolateral; B, integument; C, Al, arrow

indicates thorn-like projection.

6) P2 endopod-3 with four setae (NV. hicksi has three

and N. pectinipes has four);

7) P2 exopod-3 with only 1 outer spine (N.

pectinipes and N. hicksi with 2 outer spines);

8) P5 exopod partially fused to baseoendopod in

both sexes (not fused in N. pectinipes or N. hicksi).

The absence of an inner seta on P2 endopod-2 has

not been reported for any other member of the

Peltidiidae. Careful examination of male and female

paratypes of N. althorpensis revealed all specimens

lacked an inner seta on the P2 endodpod-2 and there

was no scar to indicate the seta had broken away.

Fusion of the P5 baseoendopod and exopod has

arisen independently in three other peltid genera:

Parapeltidium, Alteuthella A. Scott, 1909 and

Alteuthellopsis Lang, 1944 (Hicks 1986). Humes

(1981) recorded the incomplete fusion of the PS

exopod and baseoendopod in A/teuthellopsis

corallina but Hicks (1986) believed this to be an

illusion created by the orientation of the slide mount,

which disappeared when the limb was rotated. This

does not appear to be the case for N. althorpensis. All

paratypes examined exhibited partial fusion of the P5

baseoendopod and exopod.

NEW SPECIES OF NEOPELTOPSIS 18]

Fig. 8.

Neopeltopsis althorpensis sp. nov.. A, 2 Mxp

(paratype, SAM C6223), arrow indicates pad-like seta; B,

3 PI, arrow indicates ventral pectinate spine (paratype,

SAM C6224); C, 2? Pl, left arrow indicates ventral

pectinate spine, right arrow indicates patch of tiny

denticles (paratype, SAM C6223).

Based on the following apomorphic characters; the

fusion of the A2 exopod segments, the reduced size

of the Pl exopod, the reduction in setation of P2

endopod-2 and exopod-3, and the partial fusion of

the PS exopod and baseoendopod, it is suggested that

N. althorpensis is the most derived species of

Neopeltopsis (currently). Neopeltopsis althorpensis

appears to be most closely related to N. pectinipes,

sharing one apomorphic character state: A2 exopod

I-segmented. With a 2-segmented A2 exopod, N.

hicksi can be considered the least derived within the

genus; however, this species does possess an

advanced character state: 3 setae on the P2 endopod-

3 instead of 4, as found in N. althorpensis and N.

pectinipes.

Hicks (1986) suggested Neopeltopsis was most

closely related to Eupelte Claus, 1860 and

Alteuthellopsis since they all had a 2-segmented P1

endopod (see Hicks 1986; Fig. 4). Hicks (1986) also

believed Neopeltopsis and Alteuthellopsis were

related by the possession of two outer spines on the

exopod-3 of P2—P4. Strictly speaking, this character

state can no longer define the terminal clade of

Neopeltopsis and Altheuthellopsis because the P2

exopod-3 of N. a/thorpensis only has one outer spine.

However, since Neopeltopsis and Altheuthellopsis

both possess <3 spines on the P2 exopod-3 and have

two outer spines on the exopod of P3 and P4, this

clade still stands. Relationships between the other

peltid genera, as suggested by Hicks (1986), also

remain unchanged.

Neopeltopsis althorpensis is the first species of

Neopeltopsis recorded from Australia and the first

species of Peltidiidae described since Alteutha

polarsternae Dahms, 1992 was described from the

Weddell Sea (Antarctica).

In addition to N. althorpensis, five other species of

Peltidiidae were collected from Althorpe I., SA:

Peltidium simplex, Alteutha depressa, and three new

species of Alteutha, which will be described in a

future paper.

Acknowledgements

This work was supported by a grant from the

Australian Biological Resources Study. Sincere

thanks go to Alastair Hirst for collecting the

specimens and identifying the algae. I would also

like to thank the organisers of the Althorpes 2004

Expedition for providing the opportunity to conduct

this research. Thanks to Lyn Waterhouse from

Adelaide Microscopy for her assistance with the

SEM, and to the anonymous referees whose

comments helped improve the manuscript.

182 G. K. WALKER-SMITH

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REEF FISHES OF THE ALTHORPE ISLANDS AND ADJACENT COASTS

OF CENTRAL SOUTH AUSTRALIA

by S. A. SHEPHERD!, G. J. EDGAR? & N. S. BARRETT

Summary

SHEPHERD, S. A., EDGAR, G, J. & BARRETT, N. S. (2005). Reef fishes of the Althorpe Islands and adjacent coasts

of central South Australia. Trans. R. Soc. S. Aust. 129(2), 183-192, 30 November, 2005.

Reef fish populations were surveyed by visual census from 2 — 20 m depth at a range of sites at the Althorpe

Is, Haystack I., and nearby mainland reference sites. Mean densities ranged mainly from 0.13 — 0.18 m2, and

comprised mainly benthic carnivores, herbivores and omnivores. Densities are comparable to those elsewhere in

SE Australia, but much lower than those in the eastern Great Australian Bight. Forty fish species were recorded

at Althorpe Is, 28 at Haystack I. and less at the other sites. Most species were too low in abundance to examine

depth-related patterns, but the three most common species, blue-throated wrasse, herring cale and magpie perch,

decreased in abundance with depth. The main faunal differences encountered were between mainland and island

sites, and with depth. Biogeographically, the reef fish fauna was largely of widely ranging southern Australian

temperate species, but included six species with SW Australian affinities, and three species with SE Australian

affinities.

Key Worbs: Reef fishes; southern Australia; depth distributions; temperate fishes; fish abundance patterns.

Introduction

Prior to this survey, the fish assemblages of

shallow, temperate reefs of the central South

Australian coast were not formally described, and

could only be extrapolated from descriptions for the

islands of the eastern Great Australian Bight (Kuiter

1983; Branden et al. 1986; Shepherd & Brook 2003),

and from SW and SE Australia (Edgar 1984; Barrett

& Edgar 1993; Turner & Norman 1998; Curley ef al.

2002; Harman et al. 2003).

The Althorpe Islands, comprising the main

Althorpe I. and the Western Isles (see Fig. 1 in

Murray-Jones & Shepherd this issue), are situated at

the entrance of Investigator Strait, at the

juxtaposition of oceanic waters of the South

Australian Sea (Bye 1976) with waters of Spencer

and St Vincent Gulfs. Although only 8 km from

mainland Yorke Peninsula, the islands enjoy oceanic

temperatures and salinities (Bye 1976; Thomas &

Shepherd 1982), and receive nutrients from the

seasonal summer oceanic upwelling off western

Kangaroo I., which penetrates to the entrance to

Investigator Strait (Petrusevics 1993), and increases

chlorophyll a levels (Dimmlich et al. 2004). These

conditions may explain the moderately rich benthic

fauna and flora of the islands (Baker er al. 2005),

' Senior Research Fellow, South Australian Research and

Development Institute, PO Box 120 Henley Beach, S.A. 5022.

Email: shepherd.scoresby@saugov.sa.gov.au

School of Zoology, University of Tasmania, Private Bag 49,

Hobart, Tasmania. 7001.

Marine Research/Laboratories, Tasmanian Aquaculture and

Fisheries Institute, University of Tasmania, Private Bag 49,

Hobart, Tasmania. 7001.

which give the islands significance for biodiversity

conservation. Biogeographically, the fish fauna of

the Althorpe Is also has interest because the islands

lie in the transitional region between the south-

western and south-eastern Australian fish faunas

(Wilson & Allen 1987).

The Althorpe Is Expedition took place from 31

January to 10 February 2002 (Murray-Jones &

Shepherd this issue), and gave an opportunity to

survey inter alia the littoral reef fishes. Our purpose

was to describe the diversity and abundance of reef

fishes at different depths around Althorpe and

Haystack Is, and at nearby reference mainland sites,

consider the biogeographic affinities of the fish

assemblages, and assess the value of the sites as

potential marine reserves. Surveys such as this are a

vital component of this assessment process (Edgar e/

al. 1997).

Methods

Site descriptions

All shores of the islands, except within the

sheltered north-easterly facing Mooring Bay, are

subject to moderate to strong SW swells, and all are

exposed to high wind waves, with fetches variously

ranging from 12 to 100 km. The mainland sites were

exposed to the SW swell, and to wind waves from SE

to SW for Sites 8, 14 and 15, from NE to NW for Site

7, and from NW to SW for Sites 9 and 10 (see Table

1; Fig. 1, Murray-Jones & Shepherd 2005).

Rocky substratum around the Althorpe Is is

granitic, comprising mainly smoothly sloping rock

with occasional joints and caves. At Site 6 rock

184 S. A. SHEPHERD, G. J. EDGAR & N.S. BARRETT

TABLE |. Location of sites and depths sampled, with number of replicate samples (per 500 m?), at the Althorpe Is and at

adjacent mainland sites. A dashed line indicates that rock met sand above the depth shown.

Site Lat. Long. Number of replicates at each depth (m)

2 5 10 15 20

Althorpe Islands

1. The Boulders 35° 22.174' S; 136° 52.252’ E 4 5 4 3 -

2. Western Isles (NE) 35° 22.071’ S; 136° 50.952’ E l 3 4 2 l

3. Chain Islet 35° 22.448’ S; 136° 51.718’ E 0 4 4 0 0

4. Northwest Bay 35° 21.871’ S; 136° 51.406’ E 0 0 4 0 0

5. Swallowtail Bay 35° 22.480’ S; 136° 51.480’ E 4 4 4 2 2

6. Mooring Bay (W) 35° 22.011' S; 136° 51.623’ E 3 - - - -

7. Haystack Island (NE) 35° 19.406’ S; 136° 54.500’ E 0 4 0 2 -

8. Haystack I. (NW) 35° 19.266’ S; 136° 54.431’ E 0 4 0 -

Yorke Peninsula

9. Carney Point 34° 53.694’ S; 137° 00.264’ E 0 4 4 - -

10. Cape de Berg 34° 56.456’ S; 136° 58.580' E 0 4 4 - -

11. Chinamans Hat I. (S) 35° 17.419’ S; 136° 54.967' E 4 3 2 - -

12. Chinamans Hat I. (NW) 35° 17.914’ S; 136° 54.901’ E 2 - - - -

meets sand at a depth of | — 3 m, and elsewhere

around the islands at 20 — 30 m depth. At Haystack I.

and Chinamans Hat I., the substratum is travertine

limestone, and rock meets sand at 3 — 15 m according

to exposure. At Carney Point and Cape de Berg, the

substratum is granitic and rock meets sand at 12 — 15

m depth.

Transect methods

In all, 94 100 m transects were surveyed at 12 dive

sites from depths of 2—20 m at mainly 5 m intervals

(Table 1), although time constraints severely limited

sampling the deeper sites. We used a standard visual

census method (Barrett & Buxton 2002) to estimate

the size and number of all fish species within 5 m of

a 100 m or two 50 m lines set sequentially at the

given depth i.e. covering 500 m2 per transect (see

Shepherd & Brook 2003 for further details).

Experienced divers swam along the line about | m

above the algal canopy, and estimated and recorded

on a slate the length of every species. Habitats at all

sites were similar and variously dominated by the

laminarian Ecklonia radiata or species of

Cystophora, Sargassum or Acrocarpia paniculata

(Baker et al, 2005). Underwater visibility was 6 —

10 m.

Analysis

Following Shepherd & Brook (2003), reef fishes

were Classified in five ecological groups according

to behaviour and life habit as follows: Group 1 —

pelagic and mid-water species; Group 2 — species

that live in seagrasses but wander into algal habitats;

Group 3 — demersal, site-attached species that swim

above and close to the algal canopy; Group 4 —

demersal, site-attached species usually cryptic within

the algal canopy; and Group 5 — cryptic or cave-

dwelling species, active mainly at night.

To examine the relationship between fish

assemblages at the differing sites and depths

surveyed, a Bray-Curtis similarity matrix was

calculated from log (x+1) abundance data, and the

resulting site relationships plotted as two-

dimensional ordinations using multidimensional

scaling techniques (MDS) (Clarke 1993), Data were

log (x+1) transformed to avoid dominance of the

most common species, and to give greater weight to

rarer species. To avoid bias due to the differing

number of transects between sites, data from only

two transects per site were randomly selected for

analysis.

Abundances of common species were examined

for correlations with depth. Information on life habit

and nomenclature of fishes was variously extracted

from the publications of Gomon ef al. (1994), Edgar

(1997), Hutchins & Swainston (1999) and Shepherd

& Brook (2003).

Results

Assemblage patterns

The MDS plot for fish abundance data by site and

depth (Fig. 1) shows two distinct trends. First, the

Althorpe Is sites cluster together, with mainland

sites (Chinamans Hat I., Carney Point and Cape de

Berg) forming outliers. Secondly, within the

Althorpe Is cluster, fish assemblages tend to cluster

by depth for the 2, 5 and 10 m sets. We accordingly

present summaries of mean fish densities for each

depth interval for all Althorpe Is sites combined

(Table 2) and for Haystack and Chinamans Hat I.

(Table 3). Data for Carney Point and Cape de Berg

are presented for 5 and 10 m depths combined

(Table 3).

REEF FISHES OF THE ALTHORPE ISLANDS

185

fish all depths MDS

Stress: 0.23

Fig. 1. MDS ordination comparing species’ composition and abundance at 5 depths (in metres) and the 12 sites numbered

in Table 1.

TABLE 2. List of fish species recorded at the Althorpe Is at six sites from 2 — 20 m depth, ordered according to ecological

group. Mean densities for sites are given in numbers 500 m?, and mean size in cm. % occurrence is the percentage of

transects in which the species was recorded. FT is feeding type. BC = benthic carnivore; C = carnivore; H = herbivore;

O = omnivore; P = planktivore. Standard errors in brackets.

: a Density %

amieratepeeies ie 2m 5m 10 m 1S m 20m occ.

Group 1

Dinolestes lewini C 0.3 1.9 4.5 0.7 0 33

Long-finned pike (0.2) (0.9) (1.6) (0.5)

Scorpis aequipinnis P 5.3 10.2 4.0 11.3 0.3 72

Sea sweep (2.8) (4.6) (1.0) (5.9) (0.2)

Caesioperca rasor P 0 0.6 6.3 0.2 0.5 12

Barber perch (0.5) (5.7) (0.1) (0.3)

Enoplosus armatus P 0.4 1.5 3.3 0.7 1.0 60

Old wife (0.1) (0.3) (1.1) (0.1) (0.7)

Pseudocaranx dentex BC 0.1 0.1 0 0.2 0 2

Silver trevally (0.1) (0.1) (0.1)

Arripis georgiana P 0.4 0 0 0 0 2

186

TABLE 2. Cont.

S. A. SHEPHERD, G. J. EDGAR & N. S. BARRETT

: Densit %

Name:oh species rE 2m 5m 10 a 15m 20m oce.

Tommy ruff (0.3)

Mean density 6.4 14.3 18.1 13.0 1.8

Group 2

Myliobatis australis BC 0.06 0 0 0 0 2

Eagle ray (0.04)

Upeneichthys vlamingii BC 0.9 0.5 1.3 14.3 1.8 40

Red mullet (0.3) (0.2) (0.8) (1.7) (0.9)

Mean density 1.0 0.5 4.3 14.3 1.8

Group 3

Achoerodus gouldii BC 0.9 0.8 0.5 1.0 1.0 47

Western blue groper (0.4) (0.3) (0.1) (0.6) (0)

Pentaceropsis recurvirostris BC 0 0 0.05 0.2 0 4

Long-snouted boarfish (0.04) (0.1)

Dactylophora nigricans H 1.1 0.5 0.3 1.0 0.3 35

Dusky morwong (0.6) (0.3) (0.1) (0.4) (0.2)

Kyphosus sydneyanus H 0 0.2 0.1 0 0 5

Silver drummer (0.2) (0.08)

Cheilodactylus nigripes BC 6.2 3.2 2.5 1.5 2.8 84

Magpie perch (1.0) (0.5) (0.6) (0.2) (0.2)

Girella zebra H 17.9 4.3 3.4 1.7 1.0 65

Zebra fish (12.1) (2.5) (1.3) (0.8) (0.7)

Notolabrus tetricus BC 40.5 30.7 23.0 21.7 15.8 93

Blue-throated wrasse (7.9) (2.8) (3.1) (1.3) (3.4)

Dotalabrus aurantiacus BC 0.4 0.2 0.3 0 0 21

Castelnau’s wrasse (0.2) (0.2) (0.1)

Pseudolabrus parilus BC 0 0 0 1.8 0 4

Brown-spotted wrasse (1.5)

Ophthalmolepis lineolatus BC 0 0 0 0.7 0 2

Maori wrasse (0.5)

Austrolabrus maculatus BC 0 0 0.2 0 0 5

Black-spotted wrasse (0.1)

Meuschenia flavolineata oO 1.4 0.6 0.6 1.0 0 30

Yellowstripe leatherjacket (1.2) (0.2) (0.2) (0.8)

Meuschenia hippocrepis BC 0.9 0.9 0.8 1.5 0 49

Horseshoe leatherjacket (0.5) (0.2) (0.3) (1.2)

Meuschenia freycineti O 0 0.03 0 0 0 2

6-spined leatherjacket (0.02)

Meuschenia galii oO 0 0.2 0.3 0 0 16

Blue-lined leatherjacket (0.07) (0.1)

Meuschenia venusta O 0 0 0.05 0 0 2

Stars and stripes leatherjacket (0.04)

Acanthaluteres vittiger BC 0 0.08 0.10 0 0 5

Toothbrush leatherjacket (0.07) (0.05)

Eubalichthys gunnii BC 0.3 0 0 0.7 0 4

Gunn’s leatherjacket (0.3) (0.5)

Eubalichthys mosaicus C 0 0.05 0 0 0 2

Mosaic leatherjacket (0.04)

Mean density 69.8 41.6 3237 32.7 20.8

Group 4

Aplodactylus arctidens H 0 0.06 0.05 0 0 +

Southern sea carp (0.05) (0.04)

Aplodactylus westralis H 0.1 0.05 0 1.3 0 9

Western sea carp (0.1) (0.04) (1.1)

REEF FISHES OF THE ALTHORPE ISLANDS 187

TABLE 2. Cont.

: Densit %

Maung or SPevies i =m 3m Omms 15m 20m oce.

Odax acroptilus BC 0 0.1 0.1 0 0 7

Rainbow cale (0.06) (0.08)

Odax cyanomelas H 3.2 512 1.5 1.2 1.0 75

Herring cale (1.4) (1.3) (0.5) (0.8) (0)

Pictilabrus laticlavius BC 0.5 1.7 2.0 3.8 1.5 67

Senator wrasse (0.3) (0.3) (0.6) (1.7) (0.4)

Eupetrichthys angustipes BC 0 0 0.1 0 0 +

Snakeskin wrasse (0.06)

Siphonognathus beddomei BC 0 0 2.1 0 0 11

Pencil weed whiting (1.2)

Parma victoriae H 1.3 1.2 0.4 0.5 0.8 49

Victorian scalyfin (0.4) (0.4) (0.05) (0.4) (0.2)

Tilodon sexfasciatum BC 0.4 0.2 0.2 0.2 0 26

Moonlighter (0.3) (0.1) (0.1) (0.1)

Diodon nichthemerus BC 0 0.05 0 0 2

Globefish (0.04)

Mean density D5 8.5 6.5 7.0 3.3

Group 5

Parascyllium variolatum BC 0 0.06 0 0 0 2

Varied catshark (0.04)

Centroberyx lineatus P 0 0 0.05 0 0 2

Swallowtail (0.04)

Pempheris multiradiata Cc 0 4.0 6.7 0.2 1.8 26

Common bullseye (0.9) (3.4) (0.1) (1.3)

Mean density 0 4.1 6.8 0.2 1.8

Total density 82.7 69.0 65.4 67.2 29.5

Total number of species 21 28 30 23 13

TABLE 3. Mean densities of fish species (in numbers 500 m? at Haystack 1., Chinamans Hat I, Carney Point (CP) and

Cape de Berg (CdB) at various depths, ordered according to ecological group. % occurrence (the percentage of transects

in which the species was recorded) is given for Haystack 1, Chinamans Hat 1. and for Carney Point and Cape de Berg

combined. FT is feeding type. BC = benthic carnivore; C = carnivore; H = herbivore; O = omnivore; P = planktivore.

Standard errors (in brackets) are given for replicate sites or, in the case of Carney Point and Cape de Berg, for two depths.

Haystack I. Chinamans Hat I. CP CdB

Name of species FT 5m 15m % 2m 5m 10m % 5S, 5, %

Occ Occ 10m 10m Occ

Group 1

Dinolestes lewini GC 1.1 0 20 0 0 0 0 0 0.5 6

Long-finned pike (0.8) (0.4)

Scorpis aequipinnis P 12.8 10.5 80 2.9 4.4 0.5 67 10.3 12.8 69

Sea sweep (0.2) (6.4) (1.7) (0.8) (5.9) (8.0)

Scorpis georgianus P 0 0 0 0 0 0.5 7 0 0 0

Banded sweep

Enoplosus armatus P 2.0 0.5 60 0 0 0 0 1S 1.3 25

Old wife (0.4) (0.4) (1.2) (0.9)

Arripis georgiana P 5.0 0 10 0 0.2 0 7 0 0 0

Tommy ruff (3.6) (0.1)

Mugil cephalus P 0 0 0 0 0.5 0 13 0 0 0

Mullet (0.4)

Mean density 20.9 11.0 2.9: 5.1 1.0 11.8 14.6

188

TABLE 3. Cont.

S. A. SHEPHERD, G. J. EDGAR & N. S. BARRETT

Haystack I. Chinamans Hat I. GP CdB

Name of species FT 5m Sm % 2m 5m 10m % By 35 %

Occ Occ 10m 10m Occ

Group 2

Upeneichthys vlamingii BC 0.4 0 10 0 0.2 0 7 1.3 2.0 63

Red mullet (0.3) (0.1) (0.5) (0.4)

Parequula melbournensis BC 0 0 0 0 0.4 0 13 0 0 0

Silverbelly (0.3)

Siphamia cephalotes P 0 0 0 0 0 0 0 0 4.5 19

Wood's siphonfish (2.5)

Mean density 0.4 0 0 0.6 0 1.3 6.5

Group 3

Achoerodus gouldii BC 0.6 0 40 4.4 1.9 0.5 73 0.3 0 6

Western blue groper (0.3) (0.4) (1.6) (0.2)

Dactylophora nigricans H 1.3 1.0 60 1.3 1A 0 53 0.8 0.3 19

Dusky morwong (0.1) = (0) (0.9) (0.8) (0.6) (0.2)

Kyphosus sydneyanus H 2.1 0 40 4.4 3.3 0 40 12.5 75 56

Silver drummer (1.0) (0.8) (1.1) (8.2) (5.3)

Cheilodactylus nigripes BC 2.8 7.0 90 0.5 133 255: 47 3.5 5.8 81

Magpie perch (0) (2.1) (0) (0.9) (0) (1.6)

Girella zebra H 8.5 5.0 90 5.5 3.6 0 53 0.8 0.8 19

Zebra fish (0.5) (0.7) (0.4) (2.3) (0.5) (0.2)

Notolabrus tetricus BC 26.6 44.5 100 11.5 13.8 11.5 100 17.8 11.3 100

Blue-throated wrasse (4.0) (3.2) (0.4) (1.3) (2.7) (4.1)

Dotalabrus aurantiacus BC 23 3.0 80 0 0 0.5 af 0.8 0.3 19

Castelnau’s wrasse (0) (0) (0.6) (0.2)

Austrolabrus maculatus BC 0.4 0 10 0 0 0 0 1.0 0.3 19

Black-spotted wrasse (0.3) (0.7) (0.2)

Meuschenia flavolineata O 3.0 5.0 80 0 1.4 1.5 27 4.0 0.5 50

Yellowstripe leatherjacket (1.0) (2.1) (0.9) (3.2) (0.4)

Meuschenia hippocrepis BC 5.0 1.0 70 1.1 Lo: 2.0 60 9.5 4.8 75

Horseshoe leatherjacket (1.8) (1.0) (0.3) (6.3) (0.7) (2.7)

Acanthaluteres browniii BC 1.1 0 10 0 0 0 0 0 0 0

Spiny-tailed leatherjacket (0.8)

Meuschenia galii BC 0.4 0 20 0 0.2 0 7 0.3 0.3 13

Blue-lined leatherjacket (0.1) (0.1) (0.2) (0.2)

Acanthaluteres vittiger O 0.1 0 10 0.1 0 0 7 0 0.5 13

Toothbrush leatherjacket (0.1) (0.1) (0.4)

Eubalichthys bucephalus BC 0 2.0 20 0 0 0 0 0 0 0

Black reef leatherjacket (1.4)

Eubalichthys cyanoura = BC 0 0 0 0 0.2 0 7 0 0 0

Blue-tailed leatherjacket (0.1)

Chelmonops curiosus BC 0.8 0 40 0 0.1 1.5 13 0.5 0.5 13

Western talma (0.2) (0.1) (1.1) (0.3) (0.4)

Mean density 55.0 68.5 28.8 34.8 19.5 51.8 32.9

Group 4

Odax acroptilus BC 0.5 0 20 0 0 0 0 3.8 0 25

Rainbow cale (0) (2.7)

Odax cyanomelas H 10.0 1.5 100 116 7.0 1.5 100 3.5 1.5 50

Herring cale (4.1) (0.4) (0.6) (3.5) d.8) 0 (1)

Pictilabrus laticlavius BC 1.9 0 40 0.3 0.1 0.5 20 0.8 3.0 44

Senator wrasse (1.0) (0.2) = (0.1) (0.2) (1.1)

Siphonognathus beddomei BC 0.8 0 50 0 0 0 0 14.5 1.3 50

Pencil weed whiting (0.2) (8.5) (0.9)

Parma victoriae H 44 0 60 3.3 2.0 0.5 87 0.8 0.8 19

REEF FISHES OF THE ALTHORPE ISLANDS 189

TABLE 3. Cont.

Haystack I. Chinamans Hat I. CP. CdB

Name of species FT 5m 5m % 2m 5m 10m % 5; 5, %

Oce Occ 10m 10m Occ

Victorian scalyfin (1.0) (0.2) (1.1) (0.6) (0.5)

Tilodon sexfasciatum BC L.1 1.0 70 1.6 1.0 1.5 53 2.8 0.5 44

Moonlighter (0.1) (0.8) (0.3) (0.6) (0.5) (0.4)

Heteroclinus johnstoni = C 0.1 0 10 0 0 0 0 0 0 0

Johnston’s weedfish (0.1)

Mean density 18.9 2.5 16.8 10.1 4.0 26.2 7.1

Group 5

Paraplesiops meleagris — C 0.4 0.5 50 0 0 0 0 0 0 0

Western blue devil (0.1) (0.4)

Pempheris klunzingeri — C 0 0 0 0 0 0 0 0.3 1.5 13

Rough bullseye (0.2) (1.1)

Pempheris multiradiata C 1.5 1.5 60 0.8 0.7 1.0 27 0.8 2.8 25

Common bullseye (0.5) (1.1) (0.5) (0.5) (0.2) (1.2)

Mean density 1.9 2.0 0.8 0.7 1.0 1.1 4.3

Total density 97.0 84.0 49.3 513 25.5 92.2 65.4

Total number of species 28 14 14 21 13 23 24

Species richness, density and trophic groupings The abundance, Amp, of magpie perch,

In all, 40 species were recorded at the Althorpe Is,

29 species at Haystack I., 24 species each at

Chinamans Hat I. and Cape de Berg, and 23 species

at Carney Point. A comparison of Tables 2 and 3

shows that 16 species were present at offshore sites

(Althorpe Is or Haystack I.), but not at mainland

ones, and three species at mainland, but not offshore,

sites. However, these species were not abundant

anywhere, and their absence has little significance.

Mean fish densities (numbers 500 m-?) were lowest

at Chinamans Hat I. (42.0 SE 6.8) and in the range

65 — 92 for other sites, except at 20 m depth at

Althorpe I. with 29.5. The most abundant groups were

the benthic feeders (Groups 3 and 4), comprising

51% at the Althorpe Is, and variously

61 — 89% elsewhere. The second most abundant

group (Group 1) was the pelagic or mid-water species,

which comprised 12 —23% of fish numbers at all sites

except at Chinamans Hat I. where they were in low

numbers. Group 2 were in low numbers except where

transects passed near soft bottoms, and Group 5 were

present where transects crossed cryptic habitats.

Depth distributions: Three common species at

Althorpe Is showed trends of declining abundance

with depth. For the blue-throated wrasse, Notolabrus

tetricus, abundance of both juveniles (<10 cm), Aj,

and post-juveniles (>10 cm), Ajj, in numbers 500 m2,

declined with depth, D, in metres, the former

exponentially and the latter linearly. The respective

regression equations are:

Aj=5.61—2.11InD (R? =0.78; P< 0.001)

Api = 34.94 1.01 D (R2 = 0.33; P< 0.05)

Cheilodactylus nigripes, also declined exponentially

with depth, giving the regression equation:

Amp = 7.20—2.16 In D (R? = 0.33; P < 0.05)

Schools of zebra fish, Girella zebra, were

sometimes encountered at shallow depths and less

commonly with increasing depth. Due to their

schooling behaviour, variability in density was high

and the decline in abundance with depth was not

significant.

We also examined the abundance of the herring

cale, Odax cyanomelas vs mean cover of its principal

diet, Ecklonia radiata, and also vs depth. Ecklonia

cover varied between 20 and 85% (except at the

shallow western end of Mooring Bay (Site 6), where

it was virtually absent), but was not correlated with

herring cale abundance (r = 0.13; ns). If the data

from Site 6, where herring cale was also absent, are

omitted, the abundance, A,,, of herring cale declined

exponentially with depth, and the following equation

was fitted:

Ate = 9.36 - 3.05 In D (R? = 0.52; P < 0.05)

Discussion

Biogeography

The visual search method used in this study is

likely to under-estimate or overlook cryptic and

nocturnal species associated with reefs, and only by

chance includes species typical of deeper water, soft

bottoms or seagrasses. Nevertheless, the biogeo-

190 S. A. SHEPHERD, G. J. EDGAR & N. S. BARRETT

graphic affinities of the reef fauna are evident.

Wilson & Allen (1987) considered that the

southern Australian temperate region contained four

overlapping faunal components: a south-western

component, a south-eastern component, an endemic

southern component, and a broader component

extending up the east and west Australian coasts. The

Althorpe Is lie in an area of transition between the

first two components, but the reef fish fauna is

dominated overwhelmingly by widely ranging

southern Australian species. Three species

characteristic of the SW region extending to

Althorpe Is were: the western sea carp, Aplodactylus

westralis, whose eastern limit is SW Yorke Peninsula;

the spiny-tailed leatherjacket, Acanthaluteres

brownii, with eastern limits in Investigator Strait; and

the western talma, Chelmonops curiosus, with its

eastern limit in eastern S.A. Another three species,

the brown-spotted wrasse, Notolabrus parilus, the

western blue devil, Paraplesiops meleagris, and the

western blue groper, Achoerodus gouldii, extend

further east to western Victoria. Only three south-

eastern species with their western limit in

Investigator Strait or Spencer Gulf were present:

Gunn’s leatherjacket, Eubalichthys gunnii, the

southern sea carp, Aplodactylus arctidens, and

Johnston’s weedfish, Heteroclinus johnstoni. Finally,

two southern species with disjunct distributions,

absent from the colder SE Australian waters were

present: the maori wrasse, Ophthalmolepis

lineolatus, (with an eastern range now extending to

SE Tasmania — unpublished data), and the black-

spotted wrasse, Austrolabrus maculatus. None of the

above species were common at Althorpe Is.

Species abundances

Total mean densities of fish in this study (mostly

0.13 — 0.18 m~) are low in comparison with offshore

islands of the eastern Great Australian Bight, but

similar to those recorded elsewhere in SE Australia

(reviewed in Shepherd & Brook 2003). The low

densities are largely due to the low abundance of

open water feeders (Group 1), which comprise only

6 — 28% of individuals, compared with benthic

feeders (Groups 3, 4), which comprise 59 — 92%.

Hence, the ‘offshore island effect’, by which islands

capture the productivity of the surrounding ocean,

and which is reflected mainly in the increased

abundance of mid-water species, is apparently absent

from the Althorpe Is and nearby mainland sites.

Differences in terms of species composition and

abundance between sites and depths were minor, as

shown by the MDS plot (Fig. 1). We attribute this

relative homogeneity between sites to the general

topographic and habitat similarity between sites at

Althorpe Is. Algal habitats were similar around the

islands, as all sites, except those within shallow bays,

were exposed to the prevailing SW swell (Baker e7 al.

2005). The few differences that did exist can be

attributed to habitat differences (mixed rock, sand

and seagrass) and to the modified algal communities

in more sheltered sites, as in Mooring Bay, a relative

outlier in Fig. 1 (see Shepherd & Brook 2005).

Elsewhere, topographic heterogeneity, as at Haystack

L., with a high abundance of caves and crevices in an

eroded limestone substratum, favoured high fish

abundance and greater species diversity.

Although mainland sites such as Carney Point,

Cape De Berg and Chinamans Hat I. are also relative

outliers (Fig. 1), their fish assemblages overall

differed little from those of the offshore sites (Tables

2 & 3). Notable differences were: more yellow-stripe

and horseshoe leatherjackets, silver drummer, and

small blue groper (sheltered sites only), and fewer

zebra fish at mainland sites. Overall, there were no

obvious differences in habitats between Althorpe Is

and mainland sites, other than substratum at several

sites, and the obvious lack of deeper rocky habitat

(15 mand 20 m) on the Yorke Peninsula coastline due

to the shallower soft bottoms found there. The latter

difference is important, however, as, while shallow

water assemblages are similar, the presence of

assemblages and reef habitat > 40 m deep at Althorpe

Is distinguishes them from the adjacent coastline and

adds significantly to their conservation value.

With the exception of three species, referred to

below, the abundances of individual species were

generally too low to show meaningful patterns on the

depth gradient.

The three most abundant species all declined in

density with depth. This is likely due to several

factors. In the case of the wrasse, N. fetricus,

juveniles apparently recruit into shallow water, as

shown in this and the earlier study of Shepherd &

Brook (2003), and move into deeper water with

increasing size. The blue groper, A. gouldii, has the

same recruitment pattern (Shepherd 2005), but its

numbers were generally low at Althorpe Is, although

much higher densities of small groper were recorded

at sheltered mainland sites. In the case of the magpie

perch, C. nigripes, the abundance patterns may be

due to several factors. First, the majority of the 2 m

depth samples were in partly sheltered bays where

topographic complexity (abundance of granite

boulders, and crevices) was often high, compared to

deeper water. Species like C. nigripes, the scalyfin,

Parma victoriae, and wrasses, which are site-attached

and rest in shelter-holes, tend to be more abundant in

more complex habitats (Cappo 1995; Lincoln Smith

& Jones 1995; Curley et al. 2002). Thirdly, the algal

mats in which C. nigripes forages (Lowry & Cappo

1999; Wellenreuther & Connell 2002), may decline in

extent with increasing depth. Lastly, in the case of the

herring cale, the decline in abundance with depth, as

REEF FISHES OF THE ALTHORPE ISLANDS 191

at St Francis Is (Shepherd & Brook 2003), may be

associated with the general distribution of its

preferred food, the kelp Ecklonia radiata, although

we found no correlation with Ecklonia cover

measured at the scale of our sampling.

Percentage occurrence data give a different aspect

of abundance. Species with high values (see above)

are mainly either site-attached species (Groups 3, 4)

with small ranges, requiring cryptic habitat in which

to shelter, or mid-water species (Group 1), which

concentrate near vertical faces. In the case of the

bullseye (Group 5) the % occurrence value likely

reflects the frequency of large caves. Low values

indicate relative rarity.

Habitats

Curley et al. (2002) suggested that habitat-related

features were the principal determinants of the

composition and abundance of temperate fishes, and

that sites for marine parks should be selected on the

basis of conserving the maximum range of habitat

types. Topography, such as degree of relief and

substratum type (limestone or granite), depth, degree

of shelter, and algal community type are all well-

known habitat factors determining the species

present and their abundance (Harman ef al. 2003;

Lincoln Smith & Jones 1995; Shepherd & Brook

2003). There may also be benefits in conserving

island habitats, which have less terrigenous and

anthropogenic influences, but have — local

oceanographic and climatic forces, contributing to a

unique range of habitats (review of Brown &

Lomolino 2000). Habitats themselves are a good

surrogate for species, because protected areas that

incorporate the maximum number of habitat types

will also provide refuge for species, often various life

stages, and ecological linkages (Ballantine 1997;

Nowlis & Friedlander 2004). The Althorpe Is

encompass a wide depth range (including depths >15

m not found on the adjacent coastline), and a range

of exposures, giving a good representation of granitic

habitats, while Haystack I., and Chinamans Hat I.

provide a contrasting limestone topography with

high topographic complexity, and a range of

exposures and depths. Together, these sites provide a

gradient in exposure, from rough conditions at the

Althorpe Is to moderate shelter at Chinamans Hat,

and other mainland sites, and encompass a wide

range of habitats, which emir ently meet the above

criteria for marine park status.

Acknowledgements

The expedition was funded by the South Australian

Research and Development Institute and _ the

Department for Environment and Heritage. We thank

Rob Lewis, Anthony Cheshire and Bryan McDonald

for their strong support, and Sue Murray-Jones for

her organizational skill. We thank the captain and

crew of RV Ngerin, Michael Clark for manning diver

support craft, Tim Collins for providing the zodiac,

and John and Erika Lawley for their hospitality on-

shore. Chris Halstead, Ali Bloomfield, James Brook

and Renate Velzeboer assisted in the diving. We also

thank John Carragher, David Turner and Mike Steer

for helpful comments on the paper.

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Transactions of the Royal Society of S. Aust. (2005), 129(2), 193-201.

CLEANING SYMBIOSIS AMONG INSHORE FISHES AT ALTHORPE ISLAND,

SOUTH AUSTRALIA AND ELSEWHERE

by S. A. SHEPHERD!, J. TEALE? & D, MUIRHEAD?

Summary

SHEPHERD, S, A., TEALE, J, & MUIRHEAD, D, (2005). Cleaning symbiosis among inshore fishes at Althorpe Island,

South Australia and elsewhere. Trans. R. Soc, S. Aust. 129(2), 193-201, 30 Novernber, 2005.

Host-cleaner symbiosis among fish, not previously reported for South Australia, was investigated at Althorpe

I., West I., Cape Jervis and other sites over short to long periods. Five cleaner fish species, one crustacean

cleaner species, and 23 client fish species are recorded. The cleaning symbiosis is described in some detail for

two cleaners, the moonlighter, Tilodon sexfasciatus, and the pencil weed whiting, Siphonognathus beddomei, and

in less detail for three other cleaners, the western cleaner clingfish, Cochleoceps bicolor, the black-spotted

wrasse, Austrolabrus maculatus, and the old wife, Enoplosus armatus. Cleaning behaviour appears to be

facultative, and its timing dependent on cleaner behaviour. At West I. it was restricted to a few occasions per day

from late morning to late afternoon during summer months at a fixed cleaner station, whereas at Althorpe I. and

Cape Jervis cleaner stations were often not fixed, and cleaning took place opportunistically throughout the day.

A third cleaner, the western cleaner clingfish, Cochleoceps bicolor, appears to clean mainly deeper water

species. Communicative and contextual signals between client and cleaner are described. They include: guild

signs by the cleaner; posing by the client, in which the body is inclined at various angles; and fixed cleaner

stations defined variously by topography, or presence of sponges. At Althorpe I. zebra fishes that posed at an

angle head up received more attention from cleaners than those that posed horizontally, and those that posed in

small host groups received more attention than those in larger groups.

Kry Worps: Fish cleaning; cleaning symbiosis; posing behaviour; fish behaviour; ectoparasites; southern

Australia; cleaner; Tilodon sexfasciatus, Siphonognathus beddomei, Cochleoceps bicolor, Austrolabrus

maculatus, Enoplosus armatus.

Introduction

Cleaning symbiosis among fishes is a common

mutualism among tropical fishes in which cleaner

fish benefit by feeding on ectoparasites on client

species, which in turn benefit by their removal

(Losey et al. 1999). Information on the similar

temperate symbiosis is scarce, and in southern

Australia is available only from the accounts of

Hutchins (1979, 1991a) in southwestern Australia,

and brief references elsewhere (e.g. Hutchins 1991b;

Edgar 1997; Hutchins & Swainston 1999; Morrison

& Storrie 1999; Kuiter 2000). However, studies of

cleaning symbiosis in other temperate waters of the

world are more numerous e.g. the northern and

southern Atlantic (Henriques & Almada 1997;

Zander et al. 1999; Sazima ef al. 2000), the

Mediterranean (reviews of Moosleitner 1980 and

Zander et al. 1999; Zander & Nieder 1997; Zander &

Sétje 2002), the Black Sea (Darkov & Mochek

1980), the Gulf of California (McCourt & Thomson

1984), California (Hobson 1971), and New Zealand

(Ayling & Grace 1971).

' Senior Research Fellow, South Australian Research and

Development Institute, PO Box 120 Henley Beach, South

Australia 5022. Email: shepherd.scoresby@saugov.sa.gov.au

? Plant Biodiversity Centre, Hackney Rd, Hackney, South Australia

5069.

* 9 Giles St., Glenelg, South Australia 5045.

The prevalence of such cleaning behaviour among

labrids in particular has recently generated much

interest in their use as a benign method of reducing

parasite loads on fish farms (Young 1996).

Ectoparasites are a problem in southern Australian

fish farms, and observations on the symbiosis, and

the cleaner and host species involved, will be

valuable in assessing the merit of such biological

control.

In this paper we describe aspects of the ecology of

cleaning behaviour from accumulated observations

(a) at West I., South Australia (35° 37’ S, 138° 35' E)

over 15 years, (b) during the 2004 expedition and a

later visit to Althorpe I. (34° 22' S; 136° 52’ EB), (c)

at Cape Jervis (35° 35’ S; 138° 6’ E), and (d) under

Rapid Bay jetty (35° 32.8’ S; 138° 12.4’ E) with

many casual observations elsewhere.

Methods

During long-term underwater studies of abalone at

West I. (Shepherd 1998), visits were made there by

SAS at 1-2 month intervals for 1-2 days from 1983-

2000, and less often after that. Fish cleaning

behaviour was first observed in Dec. 1985 in Abalone

Cove at 4 m depth, near an abalone monitoring site, at

the junction of a vertical granite wall and a seagrass

bed of Amphibolis antarctica. Thereafter, the cleaning

station was monitored for evidence of the behaviour

194 S. A. SHEPHERD, J. TEALE & D. MUIRHEAD

up to eight times per visit at different times of day

until June 2001 when the study terminated.

Concurrently with observations of the behaviour of

the blue groper, Achoerodus gouldii (Shepherd &

Brook 2005), and the moonlighter, Tilodon

sexfasciatus, fish cleaning by the latter species was

monitored in Mooring Bay, Althorpe I. near shore at

a depth of 3 m among granitic blocks and sheet

granite. The site was monitored several times per

hour between 0900 and 1700 h from 2-11 February

and 9-13 April, 2004 and from 7-11 February 2005.

In 2005 fish cleaning was recorded mainly during

concurrent time budget studies on the moonlighter.

At Cape Jervis cleaning was observed at a

sheltered site 250 m north of Cape Jervis lighthouse

at about 3 m depth among reef of high relief with

numerous caves, in which five moonlighters resided.

At the Rapid Bay jetty site, throughout its >300 m

length, the array of vertical piles and the mass of old

piles, concrete cladding and debris strewn on the

bottom, together formed a complex reef habitat

where three cleaner species, the western cleaner

clingfish, Cochleoceps bicolor, the black-spotted

wrasse, Austrolabrus maculatus, and __ the

moonlighter, 7? sexfasciatus, were common.

Cleaning occurred at depths of 3 — 10 m at stations

adjacent to piles, among debris or near sponge.

For each cleaning event observed, data recorded

were: (a) the specific identity, number and size of the

host or client fish and cleaner fish present; (b) the

duration of the behaviour, including anticipatory

hovering or slow swimming in the vicinity; (c) pose

of the client species; and (d) the number of bites

observed for each fish. In 2004 and 2005 at Althorpe

I. we noted more carefully the angle of pose of the

client, and categorised poses as vertical (head up),

inclined (with head up at ~ 40 — 60° from horizontal),

horizontal, or nose down (at an angle of ~ 20 — 40°

from horizontal). In the horizontal category it was

not always clear whether a fish was a posing client,

or a hovering onlooker, and in cases of doubt it was

included in the latter category.

From observations on the frequency of sightings

of cleaning events at West [., an estimate of the

expected number of cleaning events there on any

day was derived as follows (see Dixon et al. 2005).

Let x be the expected mean number of daily

cleaning events during time period f, (in minutes).

If s is the total number of sightings of fish cleaning

on 7 inspections, and if the effective observation

time by the observer watching the cleaning station

is fj min. each during the period fp, then it can be

shown that:

s/n.ty X/tp

Le. x =S8.tp/ nly

The standard error of s, a binomial estimated

quantity, is given by:

V/s.(n-sy/n

Hence, from this relation, and knowing the

variance of t,, the standard error of x can be

calculated.

Note that for each observed cleaning event lasting e

minutes, if the average diver observation time is 3

min. (see below) then ¢, = e+ 6, because a positive

sighting will result if the diver observations

commence up to 3 min. before and after time period e.

Results

Five cleaner fish species, one cleaner shrimp and

23 client species, are recorded from South Australia

(Table 1). The posing position of the client fish varied

within and between species, and also according to the

cleaner species. Typically the client fish lay

motionless, dorsal side uppermost, and with fins

erect or extended. Some client species posed inclined

with head upward at various angles (e.g. Fig. 5),

while other species posed mainly horizontally (Figs

1-4). In the presence of a western cleaner clingfish or

a pencil weed whiting, the client fish often opened its

mouth and gills (Fig. 2). A summary of fish cleaning

observations at various sites, together with data on

the species involved, the duration of cleaning events,

size of client groups and number of bites per fish, is

given in Table 1, and considered in more detail below

for the main sites investigated.

West Island

At West I. fish cleaning occurred at the same station

at the junction of Amphibolis antarctica seagrass beds

and a rock wall during the 15-year period, but was not

observed elsewhere despite frequent searches.

Normally a small school of zebra fish arrived at the

cleaning station, between seagrass and a vertical wall,

and hovered in the vicinity. Client fish variously

posed horizontally, inclined or vertically, while others

swam slowly or hovered nearby. Cleaners then

emerged from the seagrass and swam about a host,

searching for and apparently picking ectoparasites

from its flanks. On four occasions other client species

were present, in addition to the zebra fish (Table 1). A

blue-throated wrasse occasionally paused nearby or

was pursued by a cleaner, but cleaning was only

observed once. On the five occasions when the

inception of cleaning was observed, the client(s)

initiated cleaning four times, by posing first and

waiting for cleaners to emerge, and on one occasion

the cleaner approached a hovering fish. All cleaning

events were recorded between December and March

between 1100 and 1500 h between 1-5 h (mean 2.4 h;

CLEANING SYMBIOSES 195

Figs 1-6. 1. Blue devil, Paraplesiops meleagris, being cleaned on the left flank by a western cleaner clingfish, Cochleoceps

bicolor. Seacliff Reef, 12 m depth. 2. Western blue groper, Achoerodus gouldii, posing for cleaning by a group of small,

almost transparent, western cleaner clingfish, C. bicolor, on the client’s flank. Western River, Kangaroo I., 16 m depth.

3. Spotted boarfish, Paristiopterus gallipavo, posing for cleaning by western cleaner clingfish, C. bicolor, which is

approaching the client near the mouth. Wirrina Reef, 22 m depth. 4. Harlequin fish, Othos dentex, being cleaned on the

flank by a western cleaner clingfish, C. bicolor. Snug Cove, Kangaroo I., 14 m depth. 5. Magpie perch, Cheilodactylus

nigripes, being cleaned on the flank by a western cleaner clingfish, C. bicolor. Note the angled pose of the client fish.

Seacliff Reef, 12 m depth. 6. Rainbow cale, Odax acroptilus, posing for a pencil weed whiting, Siphonognathus

beddomei, approaching from the rear. Note the bright eye-spot of the cleaner. Moana Reef, 10 m depth. All photos D. M.

s.e. 0.4 h) after low water.

In all there were 508 monitoring occasions

distributed across all months of the year (mean

number of occasions 42.3 (s.e. 4.8) per month), and

fish cleaning was observed only on 16 occasions,

between the months described, with sea

temperatures >16° C. We tested for independence

between the number of cleaning events and the

cumulative number of monthly monitoring

occasions for the 15-year period, and rejected the

null hypothesis of non-independence (x? = 26.6;

P<0.001) i.e. there was a highly significant

196

S. A. SHEPHERD, J. TEALE & D. MUIRHEAD

TABLE 1. Cleaner and host species with their respective length range (L), mean client abundance (A) per cleaning event,

location of cleaning, and duration, D. N = number of cleaning episodes observed. Standard errors in brackets. Nd = no

data. All sites are in South Australia, except where indicated otherwise.

Cleaner species

Client species

Pencil weed whiting

Siphonognathus beddomei

resident in seagrass

Amphibolis antarctica.

L= 5-8 cm

A=1.6 (s.e. 0.1)

Pencil weed whiting

Siphonognathus beddomei

resident in seagrass

L=5-8 cm; A=1-2.

Moonlighter

Tilodon sexfasciatus

resident in caves

L= 10-20 cm

A=1.2 (s.e. 0.1)

Moonlighter

T. sexfasciatus

resident in caves

L= 8-17 cm; A=1.

Western cleaner clingfish

Cochleoceps bicolor

L= 3 cm; A=3.

Western cleaner clingfish

C. bicolor L= 3 cm; A=1.

Resident among sponge,

ascidians and rarely bryozoans

Zebra fish Girella zebra

L=15-30 cm; A=4.1 (s.e. 0.4)

Magpie perch Cheilodactylus nigripes

L=25 cm; A=1

Bluethroated wrasse

Notolabrus tetricus

L=20-25 cm; A=1

(1) Spiny-tailed leatherjacket

Acanthaluteres brownii L=20 cm; A=1

Bluethroated wrasse N. tetricus

L=15 cm; A=1

(2) Rainbow cale Odax acroptilus

L=30 cm; A=1 (Fig. 6)

(3) Senator wrasse, Pictilabrus

laticlavius A=

Maori wrasse Ophthalmolepis

lineolata A=1

Rainbow cale Odax acroptilus A=1

Red mullet Upeneichthys vlamingii

A=1

Zebra fish G. zebra

L=20-35 cm; A=2.5 (s.e. 0.5)

Magpie perch C. nigripes

L=30-35 cm; A=1

Bluethroated wrasse N. tetricus

L=35 cm; A=1

Silver drummer Kyphosus sydneyanus

L=25 cm

Blue groper Achoerodus gouldii

L=80 cm

Old wife Enoplosus armatus L=20 cm

Moonlighter 7) sexfasciatum L=15 cm

Zebra fish G. zebra

L=20-25 cm; A=3.6 (s.e.0.7)

Magpie perch C. nigripes

L=22 cm; A=1

Blue groper A. gouldii

L=27-35 cm; A=1

Dusky morwong

Dactylophora nigricans

L=45 cm; A=1

Blue groper A. gouldii

L=30 cm; A=! (e.g. Fig. 2)

Harlequin fish Othos dentex

Magpie perch C. nigripes

L=25-30 cm (Fig. 5)

Zebra fish G. zebra L=30 cm; A=1

Harlequin fish O. dentex

L=70 cm; A=I (Fig. 4)

Blue groper A. gouldii

L=30-80 cm; A=1 (Fig. 2)

Location and notes

Abalone Cove, West I. near

vertical wall and seagrass

D=3 Isec. (s.e. 3)

Bites/fish 3.2 (s.e. 0.5)

(1)Rapid Head, (2) Moana reef,

(3) Second Valley.

((3) A.Hirst pers. comm.)

Poses generally horizontal, but

maori wrasse with inclined pose.

Mooring Bay, Althorpe I.

D=50 sec. (s.e. 9).

Bites/fish — see Table 2.

Poses of the zebra fish and

magpie perch varied (see text),

but poses of remaining

species horizontal.

Cape Jervis, 300 m north of

lighthouse, 4 m deep, near rock

wall and cave.

D=53 sec. (s.e. 15)

Bites/fish 1.5 (s.e. 0.2)

Recherche Archipelago, W.A.

Hutchins (199 1a,b)

Morrison & Storrie (1999)

Rapid Bay jetty, Western River,

K.I., Snug Cove, K.I. Second

Valley, Moana reef, Aldinga drop-

off, Hallett Cove reef, Seacliff

reef, Glenelg dredge site.

Magpie perch and zebra fish

occasionally posed inclined

CLEANING SYMBIOSES 197

TABLE |. Cleaner and host species with their respective length range (L), mean client abundance (A) per cleaning event,

location of cleaning, and duration, D. N = number of cleaning episodes observed. Standard errors in brackets. Nd = no

data. All sites are in South Australia, except where indicated otherwise (cont.).

Cleaner species

Client species

N Location and notes

Dusky morwong D. nigricans

L=50-80 cm; A=1

Boarfish Peniaceropsis recurvirostris l

L=30 cm; A=1

upward. All other species posed

horizontally.

Overall D=30-60 sec.

Spotted boarfish ]

Paristiopterus gallipavo

L=35 cm; A=I (Fig. 3)

Red mullet U. vlamingii L=25 cm; A=1 ]

Spiny tailed leatherjacket 1

Acanthaluteres brownii L=30 cm; A=1

Scaly fin Parma victoriae 1

L=15 cm; A=1

Blue devil Paraplesiops meleagris 3

L=25-30cm; A=1 (Fig. 1)

Bullseye Pempheris multiradiata |

L=15 cm; A=1

Bue groper A. gouldii 3

L=25- 35 cm; A=1

Red mullet U. vlamingii L=25cem; A=5 I

Dusky morwong D. nigricans 1

L=33 cm; A=1

Black-spotted wrasse

Austrolabrus maculatus

Rapid Bay jetty. All horizontal

poses, except zebra fish (this

study and D. Cowan (pers.

comm.))

Zebra fish G. zebra L=22 cm; A=4 1

Old wife E. armatus

L=20 cm; A=3

Cleaner shrimp

Periclimenes aesopius

Pp.

Blue groper A. gouldii L= 1.2 m; A=1 3

Leafy seadragon Phycodurus eques 1

Fam. Tetraodontidae 1

Anchorage Cove, Pearson I.

(Shepherd 2005)

Horizontal pose

Victor Harbor

Port Victoria

R.H. Kuiter (pers.comm.)

association between fish cleaning and the summer

season. Next we tested for independence between

the number of cleaning events and the cumulative

number of monthly monitoring occasions during the

above four summer months, and accepted the null

hypothesis of non-independence (y? = 4.5; ns) i.e.

the probability of cleaning occurring was

approximately the same during each month from

December to March. Next we examined the data for

time of day at which cleaning took place. There were

230 monitoring occasions at the cleaning station

between December and March, with a cumulative

mean of 23.0 (s.e. 4.1) monitoring occasions for

each hour from 0800 to 1800 h, with cleaning events

observed only between 1100 and 1500 h. The null

hypothesis of non-independence of cleaning events

and the number of observations per hour throughout

the day was rejected (x? = 29.4; P<0.001), and we

concluded that fish cleaning was restricted to the

period 1100 - 1500 h.

Lastly we calculated from the above formula that

the expected number of cleaning events per day from

December to March inclusive was 3.2 (s.e. 0.2), on

the basis that: (1) , the number of monitoring

inspections, = 132, averaging 3 min. duration each

(monitoring inspections were ~2 min. duration,

except one in four of them, which was ~6 min. during

quadrat monitoring nearby, thus averaging 3 min.);

(2) each cleaning event, including pre-cleaning

hovering, lasted 3.1 (s.e. 0.3) min., derived from five

measured events, i.e. f7 = 9.1 (see above); and (3)

both inspections and cleaning events occurred

randomly during the period 1100-1500 h.

Althorpe Island

The behaviour of the cleaner moonlighter, Tilodon

sexfasciatus, differed dramatically between 2004 and

2005 in Mooring Bay. In February 2004 three

moonlighters sheltered in a cave at the cleaning

station, and emerged from time to time to clean

groups of client species, which gathered at the cave

entrance at various times of day. Eight of the nine

cleaning events observed took place during the

period 1500-1800 h, 4-6 h after low tide (mean time

198 S. A. SHEPHERD, J. TEALE & D. MUIRHEAD

of cleaning 1558 h) and one at 1045 h at about low

tide. As previously, we tested for independence

between monitoring observations and cleaning

events, and rejected the null hypothesis (x? = 12.6; P

<0.005) of non-independence. We concluded that

cleaning behaviour strongly favoured mid- to late

afternoon. In April 2004 no cleaning took place,

despite frequent observations over a week, and some

posing by zebra fish at the cleaning station.

In 2005 several cleaner moonlighters were also

present in the same area, but sheltered in other

crevices nearby. These cleaners rarely cleaned at the

cleaning station used in the previous year, but

foraged individually or in pairs over a wide area, and

were approached or followed by client fishes in

small to large groups, seeking their attention.

Cleaning took place wherever and whenever a

moonlighter responded to a client, and was recorded

throughout the day when observations took place

(i.e. 0900 - 1700 h), and up to 5 h before and after

low tide.

Over the two years, 84% of the fishes cleaned, or

posing for cleaning, were zebra fish, while six other

species were observed posing on rare occasions

(Table 1). The zebra fish variously adopted all poses,

although the head down pose was rare, occurring on

only 1.2% of occasions. The magpie perch mainly,

and the silver drummer and blue groper always posed

horizontally, while the moonlighter posed nose down,

and the blue-throated wrasse mostly horizontally, but

sometimes inclined head up. Sea sweep, Scorpis

aequipinnis, occasionally posed, but were never

observed being cleaned, while the horseshoe

leatherjacket, Meuschenia hippocrepis, on occasions

joined a group of client species, but never posed, and

sometimes seemingly disrupted the event. Cleaners

mainly searched the flanks of a client, and

occasionally the pectoral fins, tail or head and mouth.

On only two occasions (i.e. 2.8%) did we observe

cheating, when the cleaner fish aggressively bit a

fish, causing the client to jolt noticeably and flee.

Next we examined whether attention from a

cleaner depended on the size of the client group. The

mean number of feeding bites per fish was

significantly higher for group sizes of one and two

combined, compared with larger groups (t = 3.13; P

< 0.002) (Table 2). The difference in mean number of

bites per fish between client groups of | - 2 fish and

3 or more fish was significant (t = 3.13; P < 0.002).

Moreover the proportion of fish receiving cleaner

attention declined significantly (x? = 30.6; P<0.001)

with increasing size of client group, i.e. from 87.5%

for single fish down to 25 — 45.6% for groups of 4

fish (Table 2), We then examined whether non-

horizontal posing by a client resulted in more

cleaning bites from a cleaner. Fish posing vertically

received a mean of 2.5 (s.e. 0.3) bites per fish, those

posing inclined upward received 2.0 (s.e. 0.2), those

head down received 1.8 (s.e. 0.8) while the remainder

posing or hovering horizontally received 0.5 (s.e.

0.3) bites per fish. The difference in mean bites per

fish between those posing vertically and those

inclined upward was not significant (t = 1.2; ns), but

the difference between those posing horizontally and

those non-horizontally was highly significant (t =

5.9; P<0.001). Those adopting a head-down pose

were too few to test statistically. From Table 2 it is

evident that in general 50 — 80% of fish posed

vertically or at an angle for host group sizes | — 4,

but in larger groups the incidence of posing non-

horizontally was less.

Cape Jervis

Periodic visits were made to the cleaning station

over 15 months (Table 1), but cleaning was observed

only from December to February between 1300 and

1730 h and 2-5 h after low tide. Several moonlighters

apparently resided in a cave, near the entrance to

which fishes sometimes hovered and posed for

cleaning. However, the cleaning station was very

loosely defined, as at Althorpe I. in 2005, and

cleaners often made feeding excursions away from

TABLE 2. Althorpe I. 2004 and 2005. Mean number of cleaning bites per fish for different sized groups of host fishes (all

species combined), mean percentage of fish in host group cleaned, and proportion of different posing types in each group

size category. Group Size = number of host fishes present during a cleaning event, E = number of cleaning events, and

N = total number of fish. Standard errors in brackets. For the four categories of pose, A = hovering vertically head up,

Al = inclined head up, > = hovering horizontally, and Ni = inclined head downward.

Group BE N Bites/ % Pose

Size fish cleaned ” 4) > Ny

1 32 32 1.9 (0.3) 87.5 0.16 0.37 0.41 0.06

2 17 34 1.9 (0.3) 79.4 044 O41 0.12 0.03

3 4 12 1,3 (0.5) 58.3 0.17 0.25 0.58 0

4 5 20 0.7 (0.4) 25 0.35 0.35 0.30 0

25 6 57 1.3 (0.2) 45.6 0.23 0.19 0.54 = 0.04

CLEANING SYMBIOSES 199

the cave, during which they cleaned zebra fish and

other species that sought attention.

Rapid Bay jetty and elsewhere

During the many visits to Rapid Bay jetty, cleaning

by the black-spotted wrasse and western cleaner

clingfish has been observed mainly from late

morning to mid-afternoon. The western cleaner

clingfish has been observed cleaning numerous

species at many deeper reefs (10 — 20 m) in Gulf St

Vincent and Investigator Strait where erect, stalked,

caliculate (goblet-shaped) or bilamellate sponges

(e.g. Callyspongia bilaminata), and large solitary

ascidians (e.g. Ascidia sydneiensis and Herdmania

momus) occur. The clingfish appeared to reside

within the folds, lamellae or cup, as the case was, of

the sponge, or near the ascidian, and establish a

cleaning station close by. Cleaning by the clingfish

has been observed in all months of the year, except

July and August, but this could be due to fewer diver

observations in winter months. Client species and

sites are listed in Table 1. As yet we have no data on

bite rates, due, in the case of the clingfish, to the

difficulty of obtaining data in poor light conditions,

and, in the case of the black-spotted wrasse, to the

tendency of the wrasse to retreat in the presence of a

diver.

Discussion

Most studies of cleaning in temperate regions note

that in these waters cleaning behaviour is incidental,

transient or occasional. Except for Hobson’s (1971)

study which noted a critical temperature of 12 — 13

°C, below which cleaning was rare, this is the first

study in temperate waters with evidence of

seasonality in the behaviour, presumably due to the

lower incidence of ectoparasitism in colder water

(Ayling & Grace 1971; Cété & Molloy 2003; but cf

Henriques & Almada 1997). At West I. the mean sea

temperature range 1s 13.5 °C (winter) to 21.5 °C

(summer) (Shepherd & Womersley 1970), but

cleaning was observed only from December with

temperatures >16 °C. However, if 16 °C were a

temperature threshold sensu Hobson (1971),

cleaning should occur until late May, when in fact it

was not observed after March.

Other factors, which inhibit cleaning, as noted by

Hobson (1971), include turbidity and surge. The

former is unlikely to have biased our results at West

I., where water transparency was generally higher in

winter than summer (Shepherd & Womersley 1970),

but the effect of surge or swell is not known because

monitoring observations were strongly biased toward

days with low swell and better diving conditions.

From evidence that gentle tactile stimuli are

rewarding to fish (Losey 1974; Thresher 1977),

Losey (1979) proposed that cleaning behaviour

evolved independently in many parts of the world

due to the tactile reward to the client. Cleaner

precursors exploited the positive response by clients

to such stimulation to gain ready access to a food

supply. Communicative signals evolved to increase

the probability of the response. These included:

adoption of specific poses by the client, and striking

colouration (a guild sign — see Eibl-Eiblesfeldt 1955)

by the cleaner. Contextual signals also evolved,

including cleaning stations, ectoparasite loads on the

client, and prospective food for the cleaner. In this

study some signals were present and others absent, as

might be expected from a behaviour that is transient

and occasional. Zebra fish, and occasionally magpie

perch and blue-throated wrasse, but not other

species, adopted a characteristic (non-horizontal)

pose, while other species adopted a horizontal pose

albeit with other indicia such as erect fins or opened

gills. Guild signs, in the form of conspicuous

transverse stripes, are conspicuous in the

moonlighter, the western cleaner clingfish, and the

old wife, but not the pencil weed whiting, which is a

uniform green colour. However, another possible

guild sign is the false eyespot, present in small

moonlighters, old wives, black-spotted wrasse and

female pencil weed whiting. The last species also has

very conspicuous eyes (Fig. 6), which may serve the

same function.

In this study cleaner stations, when present, had

well characterised topographies, close to rock

outcrops or vertical walls, or specific sponge or

ascidian habitats where cleaners variously resided.

However, their existence, or persistence apparently

depended on the idiosyncratic behaviour of the

cleaner species. Some moonlighters had cleaner

stations, whereas others did not. The pencil weed

whiting and western cleaner clingfish seemed to

have permanent stations. At Althorpe I. in 2004,

cleaner moonlighters adopted a ‘stay-close-to-home’

behaviour, and had fixed cleaner stations, whereas in

2005 they regularly foraged up to 30 m from their

shelter holes and cleaned opportunistically. Their

reduced foraging range may have been induced by

seal predators, which were commonly seen at the site

in 2004 but not in 2005 (cf Connell 2002).

Conspicuous topographic features characteristic of

cleaning stations were noted by Darkovy & Mochev

(1980) in the Black Sea, and by Moosleitner (1980)

and Zander & Sétje (2002) in the Mediterranean,

whereas Ayling & Grace (1971) and Hobson (1971)

found that cleaning stations at fixed sites were rare,

or, if present, loosely defined.

Diurnal peaks of cleaning activity have been

previously recorded by Johnson & Ruben (1988) and

Sazima et al. (2000), who suggested that they

corresponded with post-feeding periods of host

200 S. A. SHEPHERD, J. TEALE & D, MUIRHEAD

species, when they had time to seek a cleaning

station, At West I. schools of zebra fish often arrived

at specific points in their ranges at about the same

time of day. This may have been influenced by the

tidal cycle as cleaning was always recorded on an

incoming tide. We hypothesize that peaks of cleaning

activity are influenced by the activity periods of the

client. It is curious that at both West I. and Althorpe

I. these temporal cleaning peaks occurred only at

fixed cleaning stations, suggestive of a combined

temporal/topographic cue.

Cleaning a client’s pharynx and gills by entering

the mouth and opercular openings has been observed

only in the case of the western cleaner clingfish with

several client species (e.g. blue groper, blue devil,

dusky morwong, magpie perch, and harlequin fish).

This is not surprising as this behaviour is only

possible with a small cleaner and a relatively large

fish (cf. Ayling & Grace 1971).

The 23 client species identified in this study (Table

1) are all common in reef habitat, and are likely site-

attached but with differing range sizes. If species

with large ranges had access to more than one

cleaner fish or station (termed choosy clients), then

market theory predicts that they should have priority

of access to a cleaner fish, compared with resident

fish without such choice (Bshary & Noé 2003). At

Althorpe I., Cape Jervis and West I., zebra fish

seemed to include both residents (some were

recognizable by markings) and roving groups, but we

could not distinguish them with certainty. Hence we

could not tell whether choosy clients claimed priority

over residents. However, queuing for cleaner service

occurred when larger client groups were present, and

we never observed queue-jumping. The most

common cleaner in shallow waters, the moonlighter,

had a mean density of 4.3 (s.e. 0.9) per 2000 m? at 48

fish survey sites in southern Gulf St Vincent and

Investigator Strait (SAS unpublished data), so

potential cleaners are often spaced apart at a scale of

tens to hundreds of metres on coastal reefs. Hence

we hypothesize that these cleaners may be in

relatively short supply at many shallow coastal reefs

sites. Darkov & Mochek (1980) also believed that

there was a general shortage of cleaners in their

waters. The black-spotted wrasse (see Shepherd &

Brook 2003) and the western cleaner clingfish are

common deeper water species, but their abundance

on offshore reefs is still largely unknown

Casimir (1969), cited in Zander & Sétje (2002),

suggested that the degree of inclination in the pose of

a client species indicated its appetitive behaviour,

which is greater with steeper slope, and Coté ef al.

(1998) found that fish with an inclined posed

enjoyed twice the chance of cleaner attention

compared with non-posing fish. Others (McCourt &

Thompson 1984; Zander ef al. 1999) have reported

variable posing behaviour within a species. In this

study the 2005 Althorpe I. data suggested a more

complex picture for zebra fish. While an inclined

pose overall attracted greater cleaner attention, the

number of fishes in a client group during a cleaning

event was also an important modifying factor. For

small client group sizes of one or two, the probability

of cleaner attention was higher, and hence the

benefits of inclined posing for attention were less,

and the incidence of such poses was relatively low.

For client groups of five or more, only a minority

adopted an inclined posed, of which 96% received

cleaner attention, consistent with the hypothesis that

the rewards for inclined posing were high. We

hypothesize that inclined posing imposes an

increased risk of predation, especially at sites

frequented by seals, and hence inclined posing

mainly occurs when there is competition for cleaner

attention and the rewards of ectoparasite removal are

substantial. Our data at other sites are consistent with

this view, although of themselves inconclusive.

The low frequency (~3 — 4 events per day) and the

seasonality of cleaning, as recorded at West I., and

possibly existing at Althorpe I. and Cape Jervis,

indicate that in these temperate waters the cleaner

species involved are facultative cleaners, obtaining

only a fraction of their food from cleaning activities.

Cleaner species in other temperate seas are also

facultative (Hobson 1971; Ayling & Grace 1971;

Zander & Sétje 2002), and, according to the last-

named authors, there may be a geographic cline from

tropical to boreal waters from professional to

occasional cleaners. However, the western cleaner

clingfish may be a full-time cleaner, as at one site gut

contents were entirely parasitic crustaceans (J. B.

Hutchins pers. comm.).

Hobson (1971) suggested that cleaning stations

developed mainly at sites where fish tended to

aggregate, usually in places of high habitat diversity

ie. with some vertical relief, an observation that

accords with our experience. If cleaner species or

cleaner stations prove to be locally scarce in southern

Australia, and cleaning is a significant benefit to fish

health as suggested by some authors (Losey ef al.

1999), then their presence may provide a strong

argument for establishing reserves in their vicinity.

Many of these and related questions, such as the

ectoparasitic loads on temperate fish, fish cleaning

by deeper water species, the effectiveness of cleaning

(Grutter 1996), and the adaptive value of ectoparasite

removal in temperate waters have yet to be answered.

Acknowledgements

We thank R. K. Lewis and A. C. Cheshire for

promoting the expedition to Althorpe I., the crew of

RV Ngerin for logistic support for SAS, John and

CLEANING SYMBIOSES 201

Erika Lawley for their hospitality on Althorpe I., and

Michael Clark, James Brook, and Renate Velzeboer

for assistance in the field. Alastair Hirst, Sandra

Leigh, Steve Reynolds, Rudie Kuiter and Barry

Hutchins kindly provided data and/or advice, and

Rick McGarvey gave statistical advice. Comments

by Sue Murray-Jones, Paul Jennings and Sharon

Drabsch improved the manuscript.

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422. (Instituto Superior de Psicologia Aplicada: Lisboa).

Transactions of the Royal Society of S. Aust. (2005), 129(2), 202-208.

FORAGING ECOLOGY OF THE WESTERN BLUE GROPER,

ACHOERODUS GOULDI, AT THE ALTHORPE ISLANDS, SOUTH AUSTRALIA

by S. A. SHEPHERD’ & J. B. BROOK*

Summary

SHEPHERD, S. A. & BROOK, J. B. (2005) Foraging ecology of the western blue groper, Achoerodus gouldii, at the

Althorpe Islands, South Australia. Trans. R. Soc. S. Aust. 129(2), 202-208, 30 November, 2005.

The foraging behaviour of small to large western blue groper, Achoerodus gouldii, was examined at Althorpe I.

Small fish foraged mainly by taking feeding bites in algal canopies, or small mussels from bare rock. With

increasing size, fish switched to more efficient suction-bite feeding in which epifaunal aggregates were sucked

into the mouth, and foraged increasingly on epifaunal aggregates within algal mats. Small and medium-sized fish

tended to select foraging habitats with the highest densities of epifauna, whereas large fish, with an expanded

home range, captured large prey as well as algal epifauna. Fish at all sizes were most active, in terms of feeding

bite rate, in the morning and late afternoon, and least active in the early afternoon on an incoming tide. Small fish,

18-25 cm length, spent about half their time in shelter and foraged, swam or rested for the remainder. With

increasing size, bite rates declined, the period of emergence increased, and the proportion of time when emergent

spent swimming increased, while that spent resting decreased, and that spent foraging changed little.

Key Worbs: Western blue groper; Achoerodus gouldii; foraging selectivity; foraging behaviour; feeding mode;

time budget.

Introduction

The western blue groper (WBG), Achoerodus

gouldii (Richardson, 1843), is the largest carnivorous

fish on littoral rocky reefs in southern Australia,

reaching a length of 1.7 m (Gomon ef al. 1994). The

species ranges from the Abrolhos Is, Western

Australia to Port Phillip Bay, Victoria (Gomon ef al.

1994), but is rare on rocky reefs east of Investigator

Strait, South Australia (Shepherd & Brook'). Small

WBG are found in sheltered, shallow reef areas, and

with increasing size migrate to deeper water

(Shepherd & Brook'). The species has been targeted

by spear- and line-fishers, and its abundance

declined seriously on mainland reefs of South

Australia (Johnson 1982) until it was protected in the

Gulf waters of S.A., and its capture outside the Gulfs

made subject to size and boat limits. WBG is

proposed by the Australian Society for Fish Biology

(Crook 2001) for listing as Vulnerable (lower risk —

conservation-dependent) in the IUCN red list

category. Except for the study of Shepherd (2005)

and earlier surveys (e.g. Shepherd & Brook'), the

biology of WBG is unknown, although it appears

similar in many respects to that of its allopatric

congener, Achoerodus viridis, studied by Gillanders

& Kingsford (1993, 1998), and Gillanders (1995a,b,

*Senior Research Fellow, South Australian Research and

Development Institute, PO Box 120 Henley Beach, South

Australia 5022. Email: shepherd.scoresby@saugov.sa.gov.au

' PO Box 111, Normanville, South Australia 5204.

' Shepherd, S. A. & Brook, J. B. (2003) A survey of the western blue

groper on southern Yorke Peninsula, Reefwatch Report.

Conservation Council of South Australia. 12 pp.

1997, 1999). In that earlier study (Shepherd 2005),

the diet and foraging behaviour of WBG was

examined near Esperance and in the Investigator

Group, in the western and eastern Great Australian

Bight respectively. A common feeding mode was

shown to be suction-bite feeding in algal mats in

which the fish took epifaunal invertebrate prey non-

selectively (cf. Gillanders 1995a).

In this paper we further examine the pattern of

foraging and feeding modes employed in different

algal habitats by medium-sized to large WBG within

the framework of optimal foraging theory. The study

was carried out in Mooring Bay, Althorpe I. (34° 22’

S; 136° 52’ E), during the February 2004 expedition

there (Murray-Jones & Shepherd 2005), and during a

later visit from 9 — 13.4.04. Specifically we

addressed the questions: (1) does foraging behaviour

change with fish size; and (2) is choice of foraging

habitats influenced by prey abundance and

availability?

Methods

Site Description

The eastern part of Mooring Bay, Althorpe I. has a

smoothly sloping, granite substratum, intersected by

several shallow gullies, overlain with blocks and

boulders, and meeting sand at a depth of 3 m. The

study site extended for 135 m alongshore with an

average width of 35 m of rocky substratum, where

we followed the foraging behaviour of seven resident

medium-sized WBG, 18 — 45 cm TL, and two large

WBG, 75-80 cm TL, which foraged within and

beyond the study site.

FORAGING ECOLOGY OF THE WESTERN BLUE GROPER 203

We recognised eight distinct habitats, defined by

dominant algae or substratum features in the area,

and estimated the proportional cover of each habitat

with the line-intercept procedure. We set 100 m

transect lines parallel to shore over the length of the

study site, at 1.5 and 2.5 m depths and recorded the

distance under the line covered by each habitat type.

The transects covered the spatial extent of the range

of small to medium-sized WBG (see below). The

habitats recognised were: (1) a fucoid community,

comprising mainly, in descending order of

abundance, Cystophora monilifera, Ecklonia radiata,

Dictyota sp., C. subfarcinata, C. moniliformis, and a

sparse turf understorey of Caulerpa brownii,

Haliptilon sp. and Pachydictyon paniculatum; (2)

Caulerpa flexilis patches; (3) a coralline turf,

comprising mainly Corallina officinalis and Jania

parva; (4) a monospecific Liagora harveyana

community; (5) a monospecific P paniculatum

community; (6) cryptic habitat such as vertical faces,

overhangs or caves with a low (5-30%) cover of

Lobospira bicuspidata, Metagoniolithon stellifera,

Haliptilon sp., and Gelidium sp.; (7) bare rock; and

(8) sand. In this paper we use the term ‘turf’ to

describe structurally homogeneous, algal tufts, 3 — 5

cm high, in habitats (2), (3) and (6).

Sampling

Eight samples (each 400 cm?) in habitat (1), and

four in habitats (2) — (7), were taken by placing a

0.5 mm mesh net over the canopy/substratum,

freeing the attached algae/epifauna with a chisel, and

placing the contents in a plastic bag, which was then

sealed. Samples were preserved, and later rinsed and

weighed in the laboratory, algae identified, and

animals > 0.5 mm enumerated in the following

categories: amphipods (mostly gammarids); isopods

(including tanaids, ostracods and fragments);

shrimps (mainly Palaemonidae and Synalphidae);

crabs; gastropods; _— bivalves; _—_— polychaetes;

sipunculans; and ascidians.

Activity budget

To record the activity-time budget, four categories

of activities were recognised within three size groups

of WBG; 18 — 25 cm total length (TL), 30 — 45 cm

TL, and adults 75 — 80 cm TL. The activities

recognised (see Pavlov & Kasumyan 1998, Fulton &

Bellwood 2002) were: (a) swimming above the algal

canopy; (b) foraging, which included slow

swimming close to the algae while inspecting the

habitat, biting the substratum or epibiota, and

winnowing rejected material; (c) resting near the

bottom; and (d) interactions with other fish.

After allowing an initial 5 — 10 min. in a day for

acceptance time, a focal fish was then followed for

10 min., and the proportion of time spent in each

activity recorded during three sampling periods,

0900 — 1200 h, 1200 — 1500 h, and 1500 — 1800 h.

We counted the number of feeding bites in each

habitat, and recorded whether bites were directed

among algal fronds within the algal canopy, or on the

substratum, (which included the base of the algal

turf). Interactions with other fish were also recorded

during the same periods. The data on feeding bite

frequency were analysed by 2-way ANOVA for three

periods of day and three size-classes of fish (after

removing three 10 min. sample observations at

random from small- and medium-sized fish classes

in order to equalise replication).

To test statistically the foraging habitat preferences

of WBG we used Ivlev’s (1961) electivity

coefficient, E, which measures the degree of

selection of a particular habitat. The equation is:

E= (7% —pj)/(r; + p;)

where 7; is the proportion of feeding bites in habitat i

relative to the total number of bites, and p; is the

relative availability of that habitat, estimated by its

proportional cover within the fishes’ home range.

The index ranges from —1, for total avoidance, to +1,

for maximum selection, and values around zero

indicate random foraging. We subdivided the fucoid

habitat into two components, the canopy and

understorey, due to different foraging behaviour of

fish in each. Correlations between rankings of E and

epifaunal abundances were tested by the Spearman

rank correlation coefficient, p.

The number of emergent individuals, each

identifiable by size, was counted on 17 occasions

over 15 days at different times of day between 0900

and 1800 h. Each count was done by swimming

twice throughout the sub-adult home range, in order

to cover visually the whole area, and noting all

emergent WBG. The method gives a reasonably

precise estimate of the mean proportion of the

daytime in which fish were active, and was used by

Shepherd & Clarkson (2001) and Shepherd (2005).

Home ranges were estimated by the polygon method

i.e. by recording the maximum spatial extent

alongshore and offshore of foraging by the small-

and medium size-classes of WBG during the period

of the study, totalling > 30 h of observations.

Results

Foraging, range and habitat preferences

Small to medium-sized WBG are diurnal, benthic

feeders, intermittently resting in shelter and

emerging to feed. The smallest group (18 — 25 cm)

had a home range estimated to be 80 x 25 m ie.

2000 m? in the most sheltered part of the study site,

while the middle-sized group (30 — 45 cm) ranged

204 S. A. SHEPHERD & J. B. BROOK

0 20 40

Fish size (cm)

60 80 100

Fig. 1. Plot of the mean (+ SE) proportion, P, of feeding bites per 10-min sample directed at the substratum vs length (cm)

of western blue groper, A. gouldii, for each fish sampled. The remaining proportion of bites was directed at the algal

canopy.

further into deeper and less sheltered waters over 120

x 35 mi.e. 4200 m?*. The adult WBG foraged over at

least 15 000 m? in the Bay, and beyond its limits, but

the limited visibility of 8 — 10 m prevented us

estimating their full home range, which extended to

>15 m depth.

All WBG foraged solitarily in the various habitats

by swimming slowly ~1 — 1.5 m above the sub-

stratum, and periodically diving to the algal canopy,

turf or substratum to take one or more feeding bites.

Bites (or pecks) by small fish were directed at

individual prey in foliose algae or in algal canopies,

and with increasing size, proportionally more

feeding bites were made on the substratum. Further,

with increasing size fish more frequently employed

suction-bites, in which they closed the expanded

jaws over the algal turf close to the substratum and

sucked prey into the mouth by rapid expansion of

the buccal cavity, later ejecting the debris (see

Shepherd 2005). As it was not always possible to

detect whether fish took bites at single prey or used

suction-bites, we used data on the precise location of

bites (among algae or on the substratum), to

estimate the shift in feeding mode. A plot of the

mean proportion, P, of feeding bites on the

substratum vs fish length, L, (Fig. 1) shows the shift

in feeding location from bites among algal fronds to

bites on the substratum. The curve of best fit was the

logarithmic regression:

=-1.92+ 0.68 InL (R*= 0.96; P < 0.001)

Large fish (75 — 80 cm) foraged mainly by taking

suction-bites among algal turfs or other algal

habitats, and less often, by taking a gastropod or crab

by aggressive bite or ram-and-bite in cryptic habitat

(see Shepherd 2005).

The proportional cover of the eight habitats in the

range of small to large fish, the number of feeding

bites in each habitat, and the proportion directed at

the substratum/turf, are given in Table 1. Although

large fish (75 — 80 cm TL) foraged in deeper water

beyond the range of medium-sized fish (30 — 45 cm

TL), we used the proportional cover values for the

habitats in the home range of the latter group to

calculate habitat availability for large fish, when

foraging at the study site. None of the three size

groups foraged in the various habitats in proportion to

their availability within the fish’s range (x? = 18.4 for

the smallest size group, x? = 152.2 for the middle-

sized group, and x? =121.6 for large fish; in each case

P<0.001). For adults, we add the caveat that only the

shallow part of their range is considered here.

Values for Ivlev’s electivity coefficient, E, for the

three size groups of WBG (Table 2) indicate selected

and avoided habitats. The two smallest groups, but

not the large group, selected the coralline turf and

Caulerpa flexilis habitats for foraging. Selection for

the fucoid canopy decreased monotonically with

increasing fish size, while selection for the

substratum increased with size. The Pachydictyon

and Liagora habitats were avoided, except by the

smallest group, whereas strength of avoidance of

FORAGING ECOLOGY OF THE WESTERN BLUE GROPER 205

TABLE 1. Proportional cover of eight habitats in the respective ranges of the 18 — 25 cm and 30 — 45 cm size groups of

western blue groper, A. gouldii, and the total number of feeding bites recorded in each habitat. The proportion of feeding

bites (P,) recorded in each habitat, and the mean proportion (P.) of those feeding bites directed at the substratum for the

three size-classes of WBG are given for each habitat. Proportional cover of habitats for WBG (35 — 40 cm) applies also

to large WBG (75 — 80 cm) while foraging at the study site.

WBG (18 — 25 cm ) WBG (30 — 45 cm) WBG (75 — 80 cm)

Habitat Cover Pi P, Cover Pi, P, Ph Pe

Coralline turf 0.33 0.37 0.19 0.28 0.41 0.65 0.28 1.0

Fucoids-canopy 0.28 0.26 0 0.42 0.27 0 0.03 0

- substratum 0.02 l 0.11 I 0.44 1

Caulerpa flexilis 0.09 0.10 0.20 0.11 0.20 0.59 0.12 1.0

Pachydictyon 0.07 0.07 0.03 0.03 0.005 0 0.02 1.0

Liagora 0.11 0.14 0.06 0.08 0.002 0 0.02 1.0

Cryptic habitat 0.01 0 0 0.02 0.005 1.0 0.09 1.0

Bare rock 0.09 0.05 1 0.05 0 0 0 0

Sand 0.02 0 0 0.02 0 0 0 0

Total number of bites 478 606 129

TABLE 2. Values of Ivlev’s electivity coefficient, E, for three cryptic habitat declined with increasing size, until at

size classes of western blue groper, A. gouldii, in the largest size WBG strongly selected that habitat.

different foraging habitats in Mooring Bay, Althorpe I. Next we examined the epifaunal abundances in

Negative values indicate avoidance, and positive values algal samples from various habitats, where WBG

preference. foraged. The highest epifaunal densities were in the

Fish Size fucoid, coralline and Pachydictyon habitats with

Small Medium Large ~1800 — 3100 individuals per sample (Table 3). This

Habitat 18-25 cm__ 30-45 cm__75-80 cm was due, in the case of the fucoid habitat, to the many

Coralline turf 0.06 0.19 0 epiphytic mussels in the canopy, in the case of the

Fucoid — canopy 0.17 0.07 -0.79 coralline turf, to the mass of spionid polychaetes on

- substratum -0.90 -0.13 0.44 the substratum, and, in the case of the Pachydictvon

Caulerpa flexilis 0.05 0.29 0.04 habitat, to the many tiny rissoid gastropods on the

Pachydictyon 0 “0.71 -0.20 algal blades. Cryptic habitat and bare rock had the

Liagora 0.12 -0.95 -0.60 lowest epifaunal densities.

Cryptic habitat “I -0.60 0.64 To determine whether a relationship existed

of rock = : E between epifaunal abundance within habitats and

habitat selectivity by fish, we ranked habitats (a) by

TABLE 3. Mean abundance of epifaunal groups (numbers 400 cm”) in seven different habitats in Mooring Bay, Althorpe I.

Algal biomass is given in grams fresh weight 400 cm. Standard errors in brackets.

Coralline Fucoid habitat Caulerpa Pachy- Liagora Cryptic Bare

Species Turf Canopy Substrate —_flexilis dictyon Habitat Rock

Gastropods 134 (22) - 405 (18) 10 (2) 14093 (346) 206 (53) 2 (1) -

Bivalves 233 (68) 2900? (171) 65 (4) 150 (71) 59 (22) 1 (1) 4 (1) 54 (7)

-Amphipods 210 (11) 125 (12) 57 (9) 25 (5) 242 (57) 39 (14) 68 (4) -

Isopods - 5 (2) 24 (4) IL (2) 39 (20) 2: (1) 2 :

Shrimps - : 8 (3) - 40 (6) ‘ - L

Crabs - - 4 (1) - - - - -

Polychaetes 1423' (685) 51 (8) 59 (9) 76 (43) - - 16 (3) -

Sipunculans 18 (8) - - 2 (2) - - - -

Ascidians - - 34 (5) - - - - -

Holothurians - - 3 (1) 2 (1) - - - -

Others I - - - - - -

Total nos 2019 (782) 3081 (129) 659 (23) 276 (129) 1789 (274) 248 (67) 90 (6) 54 (7)

Biomass 203 (8) 150 (11) 43 (6) 180 (10) 108 (8) 76 (4) 27 (6) <1

‘mainly spionids. * mainly epiphytic mussels. 3 mainly rissoid gastropods.

206 S.A. SHEPHERD & J. B. BROOK

Bite rate

0900-1200

1200-1500

Time of day

1500-1800

Fig. 2. Plot of feeding bite rate (numbers 10 min."') for three size classes of western blue groper, A. gouldii, during three

time periods.

increasing value of E for each size group, and (b) by

increasing mean epifaunal density per habitat, using

data in Table 3, for each size group of fish. The

rankings were significantly correlated for small fish

(p = 0.70, P<0.025) and for medium-sized fish (p =

0.72, P<0.025), but not for large fish (p = 0.27, ns).

Hence, fish up to 45 cm TL tended to forage more in

habitats with greater epifaunal densities, a trend not

present in large fish.

Time budget

Mean feeding bite rates for each WBG size group

were relatively high during the morning, low during

the early afternoon, and high again late in the day

(Fig. 2). Bite rates were highest for the small and

middle-size groups, and lowest for the largest fish.

The ANOVA analysis showed that bite rate

differences were significant both for the fish size-

classes (F = 10.2; P<0.01) and time of day (F = 5.25;

P<0.01), with no interaction. We then repeated the

analysis for the two smallest size-classes of fish, this

time including all the data, and found no significant

difference between bites rates of the small and

medium size-classes (F = 4.07; P>0.05), although

time of day remained significant (F = 5.14; P<0.01).

In the adult group, the female bite rates were 75%

greater than the male’s, but the difference was not

significant (t = 1.1; ns). During the relevant period of

observations, the average time of low tide was 0940

h, and of high tide 1545 h. Thus the maximum

feeding rates occurred around the change of tide

shortly after low and high water respectively.

A time budget for the three size classes of WBG

(Table 4) shows ontogenetic changes in time spent

emergent, and on different activities when emergent.

However, the time budget for adults is partial only, as

TABLE 4. Activity-time budgets for three size groups of

WBG, A. gouldii, showing the percent of daytime spent

emergent, the allocation of time when emergent, to

foraging, swimming, resting and social encounters in

Mooring Bay. Standard errors in brackets.

WBG WBG WBG

(18-25 cm) (30-45 cm) (75-80 cm)

No. of 10-min 24 24 15

samples

% of total time 46.3 (6.1) 65.9 (2.6) 100

emergent

% of time emergent spent:

Foraging 49.7(9.2) 50.3 (4.0) 46.2 (7.1)

Swimming 32.1(2.1) 41.2(1.5) 51.7 (7.3)

Resting 18.1 (8.1) 8.3 (4.0) 0

Encounters:

Intraspecific 0.02 0.08 1.4

Interspecific

- Cleaning 0 0 0.7

- Aggression 0.04 0.18 -

they also foraged outside the study site, for which no

data could be obtained. The time in shelter, including

time resting during foraging excursions, declined

almost linearly with increasing size, and

concomitantly, swimming and foraging time increased

monotonically with increasing size. Intra- and inter-

specific encounters (see Shepherd 2005; Shepherd ef

al. 2005 for details) were low, indicative of a virtual

absence of competition for food (Jones 1984).

Discussion

This study provides valuable information on the

foraging ecology of a poorly known species near the

FORAGING ECOLOGY OF THE WESTERN BLUE GROPER 207

eastern end of its range. An earlier study (Shepherd

2005) has shown that the diet of WBG switched from

algal epifauna and small molluscs, in small- to

medium-sized fish, to larger fauna, such as crabs,

abalone and sea-urchins, in large fish. The feeding

mode also changed from bites or pecks at single prey

to the more efficient suction-bites on prey

aggregates, a feeding mode which became the

dominant method of feeding. The foraging data in

this paper reinforce those conclusions, as we were

able, by quantifying the precise location of bites

(algal canopy or substratum), to estimate the rate of

switching from single bites to the more efficient bite-

suction feeding mode with increasing fish size.

Feeding bites in the algal canopy or among algal

fronds are likely directed at individual crustacean

epifauna or attached molluscs, such as small mussels

or rissoid gastropods, which live diffusely among

foliose algal fronds. By comparison, bites on the

substratum are directed at the resident epifaunal

aggregates (Shepherd 2005), which live among the

accumulated detritus, rhizomes and low algal turf

(Fenwick 1976). Habitats with turfs or rhizomes

forming a low mat e.g. coralline turfs, C. flexilis, and

the substrate under fucoids, have their epifauna

concentrated within 1 ~ 2 cm of the substrate,

effectively making that habitat more profitable for

suction-feeding (Shepherd 2005). But does epifaunal

abundance in algal habitats wholly explain the

selective foraging by WBG?

While association does not logically entail

preference (Underwood er al. 2004), the strong

correlation between epifaunal abundance and E for

small and medium-sized WBG does suggest, in the

absence of other explanations, that differential prey

densities in habitats may strongly influence fishes’

choice of foraging habitat. Wellenreuther & Connell

(2002) also found for another suction-feeding

species, Cheilodactylus nigripes (see Cappo 1995), a

preference for feeding in algal mats with high prey

densities. However, other factors, such as algal

architecture, may also influence selection. Shepherd

(2005) noted that very small WBG (5 — 10 cm TL)

preferred to forage in algae with an open architecture

and lower epifaunal abundances, compared with

algae with dense fronds, and suggested that foraging

efficiency favoured a habitat that was more easily

searched visually for individual prey. This may

explain some of the noise in the data, for example

why the Caulerpa flexilis and Liagora habitats, with

lower biomass and lower frond density compared

with Pachydictvon, were favoured by some size

groups of fish. A third factor, algal patch size, was

also found by Wellenreuther & Connell (2002) to

positively affect bite rates; however, that factor

seems unlikely in this study in a between-habitat

comparison. Patches of Caulerpa flexilis, coralline

turf, and the fucoid understorey ranged in area from

0.5 — 5 m?, much smaller that the patches of

Pachydictvon and Liagora, which were mostly >10

m2? (unpublished data). Yet selectivity values, E, were

higher for the former than the latter group among the

two largest WBG size groups which employed

mainly suction feeding, suggesting that the influence

of patch size per se, if anything, was negative.

The changing pattern of habitat selection with

increasing size of fish (Table 2) is best explained by

the shift in feeding mode from single bites to

suction-feeding, which is most efficient when

directed at the substratum. The further shift by large

fish toward capture of larger prey, such as sea urchins

and abalone, with less reliance on suction feeding

(see Shepherd 2005) explains the high selectivity

value for cryptic habitat (Table 2). Omitting that

habitat and also the fucoid canopy, in which suction

feeding is ineffective, from a comparison of rankings

of E and epifaunal abundances, there is a significant

correlation (p = 0.714; P = 0.05) between the two sets

of rankings; this suggests that, in habitats where

suction feeding is employed by large fish, epifaunal

abundances influence habitat selection. Overall,

adult WBG appear to have a dual foraging strategy

for exploiting large, but sparse and patchy, prey, and

tiny, but variably abundant, prey. By foraging over a

large range, it can exploit cryptic habitats for large

prey (crabs, urchins and molluscs), and, at the same

time, take many ‘popcorn snacks’ on algal epifauna

in the ubiquitous algal habitats.

In terms of time allocation to foraging, resting and

swimming, we predict a dynamic trade-off, changing

with size, between maximising time in shelter, with

its adaptive value for predator avoidance and

reducing energy expenditure, and maximising food

intake (Hobson ef a/. 1981). Small fish are the most

vulnerable to predation and large ones need the most

food. This study exemplified these trade-offs, but

showed some differences between WBG activity at

Althorpe I., and at other sites studied by Shepherd

(2005). At Althorpe L., bite rates were lowest for the

3 h period after midday on an incoming tide, whereas

they were highest at Pearson and Ward Is for the

same period also on an incoming tide (Shepherd

2005). It is possible that diurnal activity periods are

contingent on the local ecology and unrelated to tidal

movement.

Overall, home range size, foraging preferences and

time allocation of WBG can be broadly explained by

fish size, feeding mode, epifaunal abundances and

considerations of foraging efficiency. Thus, foraging

range expands with increasing fish size and

requirement for food, habitat selection depends on

profitability, the feeding mode changes with fish size

(and a corresponding larger mouth gape) to increase

feeding efficiency, while swimming and foraging

208 S. A. SHEPHERD & J. B. BROOK

time increase accordingly. However, foraging

preferences are also dependent on a fish’s learning

and memory systems, such as spatial memory of

habitats, and memory for cues related to patch

profitabiltity within habitats (Warburton 2003), both

little known in benthic fishes. Optimal foraging

theory is likely to provide at best a heuristic and

simplified framework for studying fish foraging

behaviour in a complex environment.

Acknowledgements

We thank Rob Lewis and Anthony Cheshire of the

South Australian Research and Development

Institute for their strong support for the expedition,

and John and Erika Lawley and other friends of

Althorpe I. for their support onshore. Dive officer,

Michael Clark, provided boat and logistic support,

and the crew of RV Ngerin manned diving dinghies.

Tim Collins of Innes National Park provided a boat

during the second visit. Renate Velzeboer and Sue

Murray-Jones assisted in the field, Bob Baldock

provided algal identifications, and Janine Baker gave

much statistical assistance. Lastly we thank Simon

Bryars, John Carragher, Sue Murray-Jones and Tony

Fowler for helpful comments on the ms.

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FORAGING FLIGHTS OF SHORT-TAILED SHEARWATERS (Puffinus tenuirostris)

FROM ALTHORPE ISLAND: ASSESSING THEIR USE OF NERITIC WATERS

by L. D. Einoper!? & S. D. GoLDsworTHy?

Summary

Etnoper, L. D. & GoLpswortuy, S. D. (2005) Foraging flights of short-tailed shearwaters (Puffinus tenuirostris)

from Althorpe Island: assessing their use of neritic waters. Trans. R. Soc. S. Aust, 129(2), 209-216, 30 November,

2005.

The at-sea movement and habitat use of the short-tailed shearwater, Puffinus tenuirostris, were examined usi ng

satellite transmitters fixed dorsally to five parents provisioning chicks on Althorpe I., South Australia. Only

foraging trips of short duration were targeted by this study, and a range of flight parameters including trip

duration, foraging location, total distance covered, and maximum speed were recorded and analysed in order to

determine the behaviour of individuals undertaking short duration foraging trips. All short trips were performed

during the daytime, and lasted between 16 — 18 hours. During these trips birds foraged exclusively over neritic

waters 35 — 70 km to the southwest of the colony. Variable wind conditions did not seem to influence either the

direction of flight, or location of foraging activity. Most flight tracks were characterised by fast and direct

outbound and return flight, with concentration of time spent (putative foraging activity) in a particular area,

suggesting that birds were ‘travelling’ to a known foraging area. However, one track appeared less direct, as the

bird frequently changed direction implying that ‘searching’ flight was being performed.

Key Worbs: Short-tailed shearwater; Puffinus tenuirostris; satellite tracking: foraging; Althorpe Island; modes

of flight.

Introduction

The short-tailed shearwater or mutton-bird,

Puffinus tenuirostris, is a highly pelagic seabird, as it

spends much of its life wandering the ocean in search

of food. An individual may travel tens of thousands

of kilometres each year during an annual migration

from its breeding grounds along the southern

coastline of Australia, to the northern Pacific Ocean

(Marchant & Higgins 1990). During the austral

spring, shearwaters return to their breeding grounds,

and through summer and autumn they forage over

much of the Southern Ocean and eastern Indian

Ocean, from Australian neritic waters to the sea-ice

edge of Antarctica (Johnstone & Kerry 1976;

Nicholls e¢ al. 1998; Klomp & Schultz 2000). Much

of the research on the short-tailed shearwater has

been colony-based, and a number of long-term

studies, running for up to 55 years (Bradley ef al.

1991), have provided an understanding of their

population dynamics and breeding ecology.

Knowledge of their at-sea behaviour, however, is

very limited, due primarily to the many challenges of

recording seabird activities away from the colony. As

a result, little is known for this species about the

location of their primary foraging grounds, or the

relative importance of neritic and oceanic waters

during the breeding season. Much of our knowledge

' School of Earth and Environmental Sciences, Adelaide University,

Adelaide

? South Australian Research and Development Institute (Aquatic

Sciences), PO Box 120, Henley Beach, South Australia 5022.

of the at-sea movement and behaviour of short-tailed

shearwaters is based on observations from research

or fishing vessels (Wood 1993; Hunt et al. 1996; Ito

2002), or from the known distribution of prey items

that are returned to the colony by parents

(Weimerskirch & Cherel 1998; Connan ef al. 2005).

Their potential foraging range has also been based on

estimates of flight speed and their physiological

capabilities (Weimerskirch & Cherel 1998; Baduini

et al. 2001). However, it is now possible to monitor

the real-time location of individuals of known sex,

status and from a known colony as they travel at sea

through the use of satellite telemetry (Klomp &

Schultz 2000; Schultz & Klomp 2000a).

In the only other satellite tracking study on this

species during the breeding season, breeding adults

were recorded undertaking flights to the productive

waters of the Antarctic whilst provisioning their

chicks on their island colonies in southern New

South Wales (Klomp & Schultz 2000). These long

duration trips (10 — 30 days) are common during the

provisioning phase as adults alternate long trips with

short (I — 3 day) foraging trips in a dual foraging

strategy (Weimerskirch et a/. 1998; Schultz &

Klomp 2000b). This alternating pattern of foraging is

common among. pelagic seabird species

(Weimerskirch et al. 1994; Weimerskirch 1998;

Cartard et al. 2000; Baduini & Hyrenbach 2003;

Ropert-Coudert et al. 2004), and is thought to

represent a compromise between providing energy

flow to chicks at the nest and the maintenance of

parental body condition (Weimerskirch 1998;

Schaffer et al. 2003), Energy transfer to chicks is

210 L. D, EINODER & S. D. GOLDSWORTHY

most efficient following short trips, but at the

expense of adult condition. To restore their

condition, adults undertake longer ‘oceanic’ foraging

trips, during which chicks may fast for extended

periods (Wiemerskirch & Cherel 1998; Schultz &

Klomp 2000b). The increased frequency of feeds

associated with successive short trips gives chicks

energy for growth and body maintenance during

these fasting periods (Bradley et a/. 2000; Schultz &

Klomp 2000b). Thus, despite the short-tailed

shearwater’s exploitation of distant resources, waters

adjacent to colonies also play an important role in

breeding success.

Colony-based studies can provide information on

chick provisioning rates and the duration of parental

foraging trips, but provide little insight into the

distance travelled, path taken, or location of foraging

areas, As a result we still have a_ limited

understanding of the actual at-sea behaviour of

provisioning adults, including their general

movements, habitat use, and how their foraging

behaviour relates and adjusts to varying physical and

biological oceanographic conditions that may dictate

the location and availability of prey. This study

presents the findings of preliminary research into the

at-sea behaviour of breeding short-tailed shearwaters

from Althorpe I., South Australia (SA). Satellite

transmitters were deployed on chick-provisioning

adults, which were conducting one-day foraging trips

in order to address the following questions: (1) What

are the characteristics of short provisioning flights?

(2) When conducting short flights do adults forage

over neritic waters, or travel to nearby oceanic waters

off the continental shelf? (3) Do adults return to

specific areas to forage, or move randomly in search

of food?

Methods

Site, burrow monitoring, and adult condition

South Australia supports a population of about 2

million breeding short-tailed shearwaters distributed

over 33 breeding colonies, all located on offshore

islands (Copley 1993; Robinson ef al. 1996).

Althorpe I. is 96 ha in size, with an estimated 22,420

breeding birds (Robinson et al. 1996). The study was

carried out on Althorpe I. from 20 February —

13 March 2005. At this stage in the breeding season,

chicks are 2 — 4 weeks old (Marchant & Higgins

1990). A total of 32 burrows containing good-sized

chicks were monitored throughout the study period.

Adults that were undertaking short foraging trips

were targeted for tracking, and two methods were

used to determine the likelihood of an adult

undertaking a short trip: (1) assessing adult body

condition upon capture; and (2) monitoring the

attendance patterns of adults to chicks in the burrow.

Adult body condition was assessed by weighing the

bird to the nearest 5 g with a 200 g spring balance.

This method was used as further evidence of the type

of foraging trip that had just been completed, as it is

thought that after multiple short trips parental body

condition is low, but is increased following a long

trip (Wiemerskirch & Cherel 1998). Adults weighing

less than 600 g were deemed in low condition

(Baduini ef al. 2001), more likely to initiate a long

foraging trip (Wiemerskirch & Cherel 1998), and

were not selected for tracking. The attendance

patterns of adults were monitored by visiting

burrows daily at ~1900 h when chicks were weighed

to the nearest g with a 100 g spring balance. The

visitation of a parent could be identified by an

increase in chick weight between successive burrow

inspections. Weight increases of over 10 g were

attributed to the delivery of a meal. After monitoring

the weight changes of chicks for a period of 10 days

it was possible to identify those chicks, which had

not yet received a feed. Parents of these chicks were

deemed to have undertaken a long trip, given the

alternating foraging pattern of this species, as trips

are either 1 — 2 days, or 7 — 30 days duration

(Weimerskirch & Cherel 1998; Klomp & Schultz

2000). At 10 of these burrows, trapdoors were

installed at the entrance, with the purpose of

retaining the adult in the burrow (Weimerskirch

1998). Sticks were then placed at the entrance in

order to keep the swinging door open, and so that a

visit by an adult could be detected by the

displacement of the stick, and closure of the door,

When closed, the swinging door still allowed the

entrance of other birds, but prevented birds from

leaving the burrow. After the installation of the

trapdoors, burrow inspections were done every hour

during the night. If a visit was detected, the adult was

caught when chick begging had ceased, i.e. when

feeding had stopped.

Tracker attachment and analysis of foraging tracks

We used four 30 g KiwiSat 202 Satellite

transmitters (single AA Cell, Sirtrack Ltd, North

Havelock, New Zealand) to monitor the movements

of five short-tailed shearwaters, each weighing

600 — 700 g with a wingspan of 95 — 100 cm. Similar

sized satellite transmitters have been used

successfully on this species (Nicholls et al. 1998;

Klomp & Schultz 2000; Schultz and Klomp 2000a).

Transmitters were glued to the back feathers

(Nicholls e¢ a/. 1998; Cartard et al. 2000; Klomp &

Schultz 2000; Schultz & Klomp 2000a), using

Loctite 401 (Intek Adhesives Ltd, Northumberland,

England), and upon the birds’ return and capture at

the colony, trackers were removed. The movement of

shearwaters at sea was monitored by Service Argos

Inc. (Toulouse, France), which uses two NOAA

FORAGING OF SHORT-TAILED SHEARWATERS 211

R v

Eyre Peninsul

* Yorke Peninsula

Althorpe Is

Kangaroo Is

0 25 50

kilometers

Fig. |. The combined at-sea movements of seven foraging trips showing the exclusive foraging activity to the south west

of Althorpe I., over neritic waters. The isobaths of the continental shelf at 200 — 1000 m are shown.

(National Oceanic and Atmospheric Administration,

USA) satellites to receive a signal sent by

transmitters. The data stored on the satellites are

relayed to CNES (Centre National des Etudes

Spatiales) in Toulouse, France. The units had a

transmission interval of 60 s. Location data from

each foraging trip was analysed to determine the: (1)

location; (2) distance flown; (3) maximum distance

from colony; and (4) average travel speeds (ground

speed) between fixed locations. Tracking data were

analysed with the program ‘TimeTrack’ (version |.0-

9, M. D. Sumner, University of Tasmania, Hobart)

which filters the data by applying the filter described

by McConnell (1992) based on a maximum travel

speed of 60 km/h. We interpolated a point every 15

minutes (time) between satellite fixes, after

assuming straight-line travel between locations, and

a constant travel speed. Distance flown was

calculated as the total cumulative distance between

all positional fixes along the foraging track. For each

location point the straight-line distance from the

colony was calculated and expressed as a proportion

of the maximum distance attained during the

foraging trip. This revealed the outbound and

inbound component of each trip. The relationship

between the distance from the colony and average

horizontal speed (km h') was analysed using

regression (Spearman rank) to compare the rates of

travel during the departure and return flights from a

foraging area. Travelling speed between each point

was calculated, in order to assess changes in flight

speed during a trip and to infer two different modes

of activity. ‘Travelling’ was inferred from high speed

flight and ‘foraging’ from slow speed flight. This is

based on the rationale that birds actively exploiting a

prey patch would spend more time in a certain area

than when commuting between feeding patches

(Gremillet et al. 2004).

L. D. EINODER & S. D. GOLDSWORTHY

212

J aon 8 J ae

A Neal B Qe

eee oo ' ees Is

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ee _—

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Track 1 f Tometers Track 2 fa aomelsrs

i y = 7 J Pare

8 ¥ FA rate & & ue

\ Ln \ m

Cc Kt D ee

Q Pp @

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ys i ithorpe Is at Ry

+ a /

¢ ‘

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js 8 a 20 { a 19 20

Track 3 f ~~ Kilometers Track 4 et ~ filometers

g d J a ms As a 3 ee

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ae Alth if

bate “sg Althorpe Is q i oF my tas

\ ie av

— as

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4 Se = a

? a 724 ~~

Be out A

L § S8by ——_—— BE ed

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ot a ars

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Track 5 € ake Track 6 ra kilometers

Fig. 2. A-F The at-sea distributions of six flight paths of short-tailed shearwaters during the study period. Points represent

the output of the time-track program giving a location every 15 minutes (ie both the satellite fixes and the predicted

locations). The location of the Cape Borda weather station is shown by a star.

TABLE 1. Flight parameters from all six foraging tracks (* the same individual tracked undertaking successive trips).

Max. distance Total Average Max.

Trip Trip time from colony distance bearing speed

no. Date period (hrs) Duration (km) (km) (degrees) (km hr!)

‘? 1 25.2.05 0540-2252 17hrs12mins 70 198 232 46

is 2 26.2.05 0516-2240 17hrs 24mins 54 167 229 88.7

3 26.2.05 0410-2209 17hrs 59mins 45 143 232 53

+ 27.2.05 0356-2033 16hrs 37mins 32 130 221 44.6

5 28.2.05 0545-2154 16hrs 09mins 70 312 215 28.5

6 1.3.05 0441-2100 16hrs 19mins 37 133 240 46

FORAGING OF SHORT-TAILED SHEARWATERS 213

Wind conditions

Information on wind direction and wind strength

during the study period was obtained from the

Australian Bureau of Meteorology website for the

weather station at Cape Borda, Kangaroo I., 48 km

SSW of Althorpe I.

Results

Foraging movements

Four one-day foraging trips were monitored from

four individuals, as well as two consecutive one-day

foraging trips from a fifth individual during the study

(Table 1). Adults were away from the colony between

16 — 18 h duration. Adults began their foraging trip

either during the main departure phase from the

colony between first light (~0500 h) and sunrise

(0605 — 0609 h during the study period), or slightly

earlier at ~0400 h, as fewer adults chose to depart

before first light. All birds travelled over neritic

waters to the SW of Althorpe I. (Table 1; Figs 1, 2A-

F). The maximum distance from the colony ranged

from 32 — 70 km, and the total distance travelled

during foraging trips ranged between 130 — 312 km

(Table 1).

The proportion of maximum distance was

negatively correlated with the average horizontal

speed on both the outbound (Spearman’s rho: r =

0.563, P = 0.01) and return (Spearman’s rho: r =

0.172, P = 0.05) foraging trips, indicating that

shearwaters travelled more rapidly when they were

close to the colony, compared with when they were

close to their maximum distance (Fig. 3A). This

result is due to the similar foraging behaviour

performed during five of the six tracks, involving

‘travelling’ movement to an area followed by a period

of ‘foraging’, before returning to the colony (Figs

2A-D, and F, and Fig. 3B). In contrast, Track 5

showed a different pattern of movement, undergoing

many changes in direction (Fig. 2E), more variable

flight speeds throughout the trip duration (Fig. 3B),

reaching maximum distance towards the end of the

trip (Fig. 3A), and covering a greater total distance

(Table 1). These features are indicative of ‘searching’

flight (Arnould e¢ al. 1996; Weimerskirch ef al.

1997), and suggest that the individual was actively

searching for a suitable patch of prey. The location of

this individual’s movements also differed from that

of the four other individuals involved in the study.

The short period of ‘foraging’ flight was located

close to the coastline of Kangaroo I., whereas the

other birds foraged further from the Kangaroo I.

coast, a few km to the north and west.

Relationship with wind conditions

The wind strength was moderate to strong during

the study period, varying from 11 — 33 km hr! (Table

Distance from colony [%]

0-10 10-20 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100

Stage of foraging trip (+)

Flight speed ({kmfh]}

0-10 10-20 20-30 30-40 40-50 50-60 60-70 70-80 80-90 90-100

Stage of foraging trip (+)

Fig. 3, Relationship between (A) the average distance from

Althorpe I., as a proportion of maximum distance (%),

and (B) the average horizontal speed (km hr) travelled

between 15 minute points, at different stages of the

foraging trip (% duration). Vertical bars are standard

errors. Solid circles show the mean for Trips 1, 2, 3, 4,

and 6 and open circles represent Trip 5.

TABLE 2. Wind direction and speed (km Ir!) at Cape Borda,

Kangaroo 1.

Date Time Direction Speed

25.02.05 900 SE 11

1500 SW 31

26.02.05 900 ESE 15

1500 N 19

27.02.05 900 NE 24

1500 NNE 17

28.02.05 900 NE 19

1500 WSW 13

1.03.05 900 S 30

1500 S 33

2). Wind direction was variable during the five

consecutive days of the study. The fact that all

foraging trips were directed to the SW of Althorpe I.

indicates that neither the strength nor direction of the

wind had a measured influence on the flight

behaviour of tracked birds.

214 L. D. EINODER & S. D. GOLDSWORTHY

Effect of transmitters on shearwaters

The duration of the foraging trips of tracked

individuals, as well as their successful provisioning

of chicks upon return to the colony, suggest that the

attachment of trackers had minimal adverse effects

on foraging behaviour. In addition, tracked birds

departed and returned to the colony with the main

movement of the rest of the colony.

Discussion

Habitat utilisation

These results represent the most detailed tracking

data for the short duration foraging trips of this

species, and provide an insight into their foraging

ecology over the neritic waters surrounding their

colony. This study revealed that adults concentrated

their foraging efforts over a body of water 30 — 70

km SW of the colony in the week of the study. A

number of studies on other pelagic seabirds have

revealed that tracking data can be used to draw

inferences as to the location and predictability of

patches of prey in the marine environment (Anderson

et al. 1997; Weimerskirch et al. 1997; Ropert-

Coudert et al. 2004). The concentration of foraging

effort by five individuals to the SW of the colony

suggests that adults were not foraging randomly from

the colony. Individuals may be re-visiting a known

prey patch exploited the day before, either on their

previous short trip, or on their return from a long trip.

The bird that was tracked undergoing two short trips

to the same area exemplified this notion of an

individual returning to an area of successful foraging

over successive day trips.

The flight capabilities of the short-tailed

shearwater (Klomp & Schultz 2000; Baduini er al.

2001) reveal that provisioning adults are quite

capable of conducting a 200 km round trip to the

edge of the continental shelf, or beyond, in the 16-18

hours they are absent from their colony.

Nevertheless, this study suggests that during

foraging trips of short duration they are closely

linked to shelf areas of less that 100 m depth. Only

one other tracking study has recorded the short trip

movements of this species during the chick-

provisioning phase (Schultz & Klomp 2000b). That

study, conducted of a colony in NSW, recorded two

successive day trips of a single breeding adult, and

revealed that foraging was restricted to the strip of

neritic water south of the colony. The foraging

parameters including flight speed and distance

covered were comparable to this study.

Fisheries interactions

The results of this study show that breeding adults

from Althorpe L. concentrated their foraging effort in

an area close to that utilised by the SA pilchard

(Sardinops sagax) fishery (Rogers et al. 2004). The

role of pilchards in the diet of the shearwater

population in SA is unclear because no dietary

studies have been conducted in this region. However,

shearwaters are known to feed on small pelagic

fish species in other parts of their breeding range

to the east (Montague et al. 1986; Skira 1986;

Weimerskirch & Cherel 1998). Foraging location and

dietary data for short-tailed shearwaters are needed

in order to determine the degree of spatial overlap

and competition between shearwaters and

commercial fishers.

Distribution of food resources

Many seabird species have proven to be a useful

tool in monitoring the marine environment, because

fluctuations in seabird population, breeding, or

nutritional parameters are commonly due to

perturbations lower in the food chain (Cairns 1992;

Croxall et al. 1988). Foraging parameters are

particularly sensitive to changes in prey availability

(Cairns 1992; Montevecchi 1993), and thus can.be of

value as indicators of changes in the quality,

abundance or availability of prey. The link between

shearwater foraging behaviour over neritic waters of

Australia, and the availability of marine food

resources is not well understood. Despite a small

sample size, the area of concentrated foraging

activity to the SW of Althorpe I. suggests that this

body of water supported adequately profitable prey

patches during the weeklong study period. The

replication of these tracking methods over a number

of successive weeks or months during the chick-

provisioning phase would enable further assessment

of the dynamics of prey availability over neritic

waters surrounding Althorpe I.

Oceanographic features

There is growing evidence that marine birds do not

randomly sample the environment for prey, but focus

their search where prey can be located predictably.

Spatially and temporally predictable feeding sites,

such as those associated with upwelling, are of

importance to breeding Procellariiformes that

depend upon the availability of prey near their

colonies in order to raise young (Ainley ef al. 1998).

A series of regional coastal upwelling centres occur

along the SA coastline, driven by SE winds in

summer (Kampf ef al. 2004); however, the spatial

and temporal variation in upwelling intensity in the

eastern Great Australian Bight has only recently

received attention, and the reliance of many local

apex predator populations on the productivity

associated with these areas is little known (Gill

2002). The shearwater population on Althorpe I. is

well located to exploit the marine productivity

associated with these events, as a site of strong

FORAGING OF SHORT-TAILED SHEARWATERS 215

upwelling is located along the western coastline of

Kangaroo I., between 60 — 90 km away (McClatchie

et al. 2005). The concentration of foraging effort to

the SW of the colony reported in this study may have

been in response to the increased marine productivity

associated with upwelling activity in that area;

however, the strength of this relationship is reliant

upon an assessment of the wind conditions, sea

surface temperatures, and data on the primary and

secondary productivity in the area during the week of

the study. Future research into this topic requires the

co-ordination of oceanographic studies in the area,

with tracking and dietary studies of this species, on

the same spatial and temporal scales.

Conclusions

This study provides the first information on the at-

sea behaviour of shearwaters when undertaking short

foraging trips over neritic waters in the eastern Great

Australian Bight. Due to the co-ordinated departure

and return of tracked birds with the rest of the colony,

and their successful provisioning of chicks, it

appears that 30 g satellite tracking devices are a

feasible method of monitoring the normal at-sea

behaviour of provisioning adults. Most tracked

foraging flights clearly revealed two modes of travel:

direct ‘travelling’ flight to a particular area, and

slower flight associated with ‘foraging’ activity. This

enabled the identification of a concentrated foraging

area over neritic waters to the SW of the colony. The

consistencies in flight direction from the colony

suggest that birds do not forage randomly, and that

foraging flights are undertaken towards a specific,

possibly previously known, area.

As the short-tailed shearwater is one of the most

abundant pelagic seabirds over the neritic waters of SA

in summer, they must consume a large amount of the

area’s marine resources. Of primary concern is their

interaction with the harvest of pilchards (Sardinops

sagax) by commercial fishers. Such competition is of

growing interest to environmental and fisheries’

managers in many parts of the world (Tasker et al.

2000; Goldsworthy ef al. 2001; Furness 2003). As

fishing effort is concentrated 50-100 km west of

Althorpe I. (Rogers et al. 2004) and may expand to

other areas in future years, the potential exists for

competition between shearwaters and commercial

fishers. Satellite tracking and dietary work is pivotal to

achieving an understanding of such competition.

Acknowledgements

We thank those who assisted in the collection of

data whilst in the field, including Brad Page, James

Thiessen, Sarah Pennington, Deb Fraser, and Jon

Pipitone. To John Lawley for a guided tour of

Althorpe I., and for his interest and enthusiasm in the

research project. Thanks also to Tim Collins and the

staff of Innes National Park for the loan of

equipment, and assistance in transport when in the

field. The program ‘Timetrack’ was designed by

Mike Sumner, of the University of Tasmania, and

kindly provided for the analysis and filtering of the

tracking data. We also thank referees and editors for

improvements to the manuscript.

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