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PROCEEDINGS

OF THE

LINNEAN SOCIETY

NEW SOUTH WALES

VOLUME
111
(Nos 485, 486, 487 & 488; for 1989)

Sydney
The Linnean Society of New South Wales
1989

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Index

Volume 111

Page
Acacia iteaphylla, seedling development .... 37
Acanthodians, culmacanthid ........... 11
Achaearanea mundula ................... 25
Amino Acids, Amides, redistribution .... 37
Amphientomidae, two new species ...... 31
Anderson, G. J., seeGray, M. R......... 25
Antarctica, south eastern Australia ...... 11
ANMIECIOIONGAS 5.5 0000050000000000000008 278
Araneoidea, Theridiidae .............. 25
Argyrodes, host predation ............... 25
Argyrodes incursus sp. NOV. ............--- 25
Asteroidea, anew genus............... 293
/ASURODSCUUONGIENS © oo onc 0e 9050040040408 268
Austrochthonius australis, redescription ..... 233
Belyaevostella gen. nov. ................. 304
Belyaevostella hispida ................... 304
TBST REGIS 5 coc occossec0esesncns 67-100
Bradke, A. B., & Murray, D. R., Redistri-
bution of amino acids and amides
during seedling development in Acacia
iteaphylla F¥. Muell. (Fabaceae: Mimo-

SOIGEAS) Mea rey sorte jens wiser ores OR eee 37
Brasimerdaes 92 bine We eae Sa eee 274
Brown, Robert, Australia 1801-5 ........ 65
Bry opliyitayen cic ssc nee ae one 82
Buchanan, R. A., Pied currawongs (Strepera

graculina): their diet and role in weed

dispersal in suburban Sydney, New

SouthsWales so oe ccs sd vee eee 241
Caenopedina alanbakert sp. nov ........... 265
Campbell, H., John Vaughan Thompson,

[By Lie Sis, eeaue tears tie ree Cee Nace nee nerece 45
Caymanostella admiranda ................ 298
Caymanostella Belyaeva, new species ==
GESCHIPUOM Sascha oe aes ome ctne yer: 293
Caymanostella phorcynis sp. nov. .........-. 301
Caymanostella spinimarginata ............. 298
Caymanostellidae, a review ............ 294
@lhiarophyitai pac qe5 6 ces ans cae sete eee we 77
Chernetidae, new Australian species ..... 123
Chlorophyta fie .sshqacnessncsansess 70
Chthoniidae (Pseudoscorpionida) ....... 233
Gurcipedesa acoso Are aacia wamern cin cee 53
Conicochernes doyleae sp. nOv ...........-. 123
(COMMERO NSS . 600065050000 00n00005 96
Cryptogams, Gymnosperms ........... 65
Culmacanthus antarctica sp. nov ........... 14
Culmacanthus pambulensis sp. nov ......... 17
(CHYCAGCIOD VAIS conn cccccccasccanenonge 96
IDsyormein, IPISEES .5200000c0c0a0n050000 11
Diet and role in weed dispersal in suburban

Swieleyg Air soca iyi ianyee SEN ORS Se rea 241

Echinoderm fauna checklist ............
Echinodermata, deep-water species from
Norfolk Island and Wanganella Bank .
Echinothunidacinn nnn panononerorncer
Elix, J. A., & Streimann, H., The Lichens
of Norfolk Island. 1: Introduction and
the Family Parmeliaceae............

Fabaceae, Mimosoideae...............
Filicopsida (True Ferns) ...............
Flannery, T. F., Microhydromys musseri n. sp.,
a new murid (Mammalia) from the
Torricelli. Mountains, Papua New
Guiineare cei: oo renee nc ee
JNO DARD 20262 000c000c0000000000¢
Flavoparmelia norfolkensis sp. nov. .........

Glyphodiscus mcknighti ................4.
GOMASISHCES, oocccacceaccc00000008
Gray, M. R., & Anderson, G. J., A new
Australian species of A7gyrodes Simon
(Araneoidea: Theridiidae) which preys
ONES MOSE psagis sce race aeeahatenaeaytiecee lo
Groves, E. W., & Moore, D. T., A list of the
cryptogams and gymnospermous plant
specimens in the British Museum
(Natural History) gathered by Robert
Brown in Australia 1801-5 ...........

Hapalosoma pulchrum sp. nov. ........-.--
Hemuiseopsis alettae sp. nov. ............-.
History, Murrary Cod Fishery ..........
Holothuria (Vaneyothuria) unica sp. nov .....
Iolothuriid cae neeeeeeeeeceeeeeeneer

Kennedy, C. M. A., Redescription of Aus-
trochthonius australis Hoff (Chthoniidae:
Pseudoscorpionida) ...............

Kennedy, C. M. A., Conicochernes doyleae, a
new Australian species of the Cherneti-
dae (Pseudoscorpionida: Arachnida) .

Lake Macquarie, New South Wales .....
Lambert, M. J., & Turner, J., Redistri-

bution of nutrients in subtropical rain-

forest trees! 2. sion a Sees ae
Larval silverbiddy Gerres ovatus and Gobies
Weichemes: 2c sented sya eens sees ae ae
Liverworts, leafy, thalloid..............
Lophostemon confertus .............-.-.--
Lycopsida “<5 ssacs adasgene an sabes acne

Maccullochella peeli .................4..
Mckenziartia, Pectocythere................
WMickenzvantiah) qucctae eee ee

257

263

103

37
85

215
110
110

273
273

233

123

PROC. LINN. SOC. N.S.W., 111(4), 1989

Mckenziartia mowbrayi sp. nov. ...........
Mckenziartia portjacksonensis .............
Mckenziartia thomi sp. nov. ............-.
Mesothuria (Pentchrothuria) norfolkensis sp. nov
Mucrohydromys mussert sp. NOV. ...........
Mikulandra, M., see Yassini,I...........
Moore, D. T., see Groves, E.W. .........
Murray, D. R., see Bradke, A. B.........
Murid (Mammalia), Torricelli Mountains
IMNISClis.rc cea nerd yee eas HeReRSEO kor Rae ae

Nanometra duala sp. nov. ................
Neo fuscelea® inc. kssiis. wie ays 1 st Ws eSewackten wars tie
Neofuscelia verrucella ...................
Neothyonidium parvipedum sp. nov. ........
Nitrogen, mobilization ................
Norfolk Island, lichens ................
Norfolk Island, Wanganella Bank, north-

eastern Tasman Sea ...............
Novodiniainelenach nee eee ee
Nutrients, redistribution ..............

Ordovician — Silurian Stratigraphy .....

Parapanmelia A. 4.2 pent eho = ee
Paraparmelia scotophylla .................
Rarmeliag yt. tse d canes: € ona boeaine
MALO. GOUMIVAIS x0008000000000000000¢
Parmeliaceae, family .................
Rarmelinopsis viscevsteer ayo kes et ener:
Parmelinopsis spumosa .............+4-.-
IParmotremal ae Ak Sustains Hae hs SR
Parmotrema austrocetratum ...............
Parmotrema chinense .................-..
Parmotrema crinitum ....................
Parmotrema cristiferum ..............++5.
Parmotrema gardnert ....................
Parmotrema rampoddense .................
Parmotrema reticulatum...................
Parmotrema sancti-angelit ................
Parmotrema tinctorum .............+++.--.
Pectocythere royi sp. nov. ..............-.-
Pectocytheridae, Ostracoda, Crustacea ..
Pedinidaers-1e) sere to eae re
Pemberton, J. W., The Ordovician —

Silurian stratigraphy of the Cudgegong

— Mudgee District, New South Wales
Rentacnimusscuropacus ene ee Eee
Percichthyidaeneae eee ee
Phacophytacastiei Moe oe ee eee ee See
Pied Currawongs (Strepera graculina) ......
Phyllophonicdaceen eee eenneraereeee
iRiscess Devonian) eee eno
ROLY 20a: oro ther it ee eee
Pressey, R. L., Wetlands of the lower

Clarence Floodplain, northern coastal

INERT SOWEN WANES 50c00cc0cg0cd00c0
Pressey, R. L., Wetlands of the lower

Macleay Floodplain, northern coastal

New South Wales .................

PROC. LINN. SOC. N.S.W., 111(4), 1989

143

157

INDEX

Pseudoscorpionida, Arachnida .........

Psilopsidad i. < snc cers one sce ee one
Psocoptera, Insecta ...................
Pteridophyta: <0.) son oanusee Gea es
Rainforest trees, subtropical ...........
Rhodophyta’ <:¢4.5..24.52.25 See ee
Rowe, F. W. E., A review of the family

Caymanostellidae (Echinodermata:

Asteroidea) with the description of a
new species of Caymanostella Belyaev
and a new genus ...................
Rowe, F. W. E., Nine new, deep-water
species of Echinodermata from Norfolk
Island and Wangangella Bank, north-
eastern Tasman Sea, with a checklist of
the echinoderm fauna..............
Rowland, S. J., Aspects of the history and
fishery of the Murray Cod Maccul-
lochella peels (Mitchell) (Percichthyidae)

Seopsis incisa sp. NOV. ........-++-+--+-0
SiltstonesyAztee Sam aa sane eee cee
Smithers, C. N., Two new species of

Amphientomidae (Insecta: Psocop-
tera), the first record of the family for
Australia: fcc ccsteccius ten sree

Steffe, A. S., Tidal and diel variations in
the abundance of larval fishes in Botany
Bay, New South Wales, with emphasis
on larval silverbiddy Gerres ovatus
(Fam. Gerreidae) and gobies (Fam.
Gobiidae) ee. Cae ee eae

Stratigraphy, Cudgegong-Mudgee District

Streimann, H., see Elix, J. A. ...........

Synallactidae: . 2.45. JE. a

Tethyaster tangaroae sp. nov. ..............
Theridiidae, Araneoidea ..............
Turner, J., see Lambert, M.J............

Variations in abundance, tidal, diel,
Botany Bay) oo. cretistameatsteeoe

Weed dispersal, currawongs ............
Wetlands lower Clarence floodplain .....
Wetlands lower Macleay floodplain ......

Xanthopanneliaaeeen eee er ee
Xanthoparmelia amplexula ...............
Xanthoparmelia australasica ..............

Yassini, I., & Mikulandra, M., Mckenziartia
and Fectocythere (Pectocytheridae, Ostra-
coda, Crustacea) in Lake Macquarie,
New South Wales .................

Young, G. C., New occurrences of
culmacanthid acanthodians (Pisces,
Devonian) from Antarctica and south-
eastern Australia ..................

293

131

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 111
NUMBERS 1-4

NATURAL HISTORY IN ALL ITS BRANCHES

THE LINNEAN SOCIETY OF
NEW SOUTH WALES

Founded 1874. Incorporated 1884.

The Society exists to promote ‘the Cultivation and Study
of the Science of Natural History in all its Branches’. It
holds meetings and field excursions, offers annually a
Linnean Macleay Fellowship for research, contributes to
the stipend of the Linnean Macleay Lecturer in Micro-
biology at the University of Sydney, and publishes the
Proceedings. Meetings include that for the Sir William
Macleay Memorial Lecture, delivered biennially by a
person eminent in some branch of Natural Science.

Membership enquiries should be addressed in the first
instance to the Secretary. Candidates for election to the
Society must be recommended by two members. The
present annual subscription is $35.00.

The current rate of subscription to the Proceedings for non-members is set at $50.00 per volume.

Back issues of all but a few volumes and parts of the Proceedings are available for purchase. A price list will
be supplied on application to the Secretary.

OFFICERS AND COUNCIL 1988-89

President: T. G. VALLANCE

Vice-presidents: P. M. MARTIN, HELENE A. MARTIN, C. N. SMITHERS,

Honorary Treasurer: 1. G. PERCIVAL

Secretary) BARBARA J. STODDARD

Council: A. E. J. ANDREWS, T. C. CHAMBERS, JUDITH H. K. EASTMAN,
M. R. GRAY, SUSAN J. HAND, D. S. HORNING, L. A. S. JOHNSON,
R. J. KING, HELENE A: MARTIN, P. M. MARTIN, J. Ri: MERRICK,
2a IMINO SCOUT sl SG MEINE | IPO IDL, vhs UNINC S08.
C. N. SMITHERS, T: G. VALLANCE, KAREN L. WILSON

Honorary Editors: T. G. VALLANCE — Department of Geology & Geophysics,
‘University of Sydney, Australia, 2006. (Numbers 1 & 2.)
J. R. MERRICK — Graduate School of the Environment, Macquarie
University, North Ryde, N.S.W., Australia, 2109. (Numbers 3 & 4.)

Linnean Macleay Fellow: JULIA M. HUSH

Linnean Macleay Lecturer in Microbiology: K.Y. CHO

Auditors: W. SINCLAIR & CO.

The postal address of the Society is PO. Box 457, Milson’s Point 2061, N.S.W.,
Australia. Telephone (02) 929 0253.

Cover motif: The gastropod collected ‘16 miles cast of
Wollongong’ and described by Charles
Hedley as Stiva ferruginea gen. ct sp. nov.
(Proc. Linn. Soc. N.S.W. 29, 1904: pl. LX,
no. 23).

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 111
NUMBER 1

Redistribution of Nutrients in Subtropical
Rainforest ‘Irees

MARCIA J. LAMBERT and JOHN TURNER

(Communicated by D. W. EDWARDS)

LAMBERT, M. J., & TURNER, J. Redistribution of nutrients in subtropical rainforest
trees. Proc. Linn. Soc. N.S.W. 111 (1), 1989: 1-10.

Fresh foliage and leaf litterfall from trees and understorey plants in a N.S.W. sub-
tropical rainforest were chemically analysed to estimate nutrient redistribution. In
general, the proportion of nutrients redistributed in these species at time of leaf litterfall
is low. Results available on nutrient redistribution from the same species in this area
during heartwood formation showed that trees which redistributed phosphorus from
foliage, redistributed little from heartwood and vice-versa. By way of contrast, species in
sclerophyll forests were highly efficient at nutrient redistribution from both leaves and
wood. Oo me peters

Marcia J. Lambert and John Turner, Forestry Commission of N.S.W., PO) Bil aomh BeeBibl datcal | air
tralia 2119; manuscript receved 19 November 1986, accepted for publication 23 TES eG
f.

| IBRARY
INTRODUCTION j 1
Nutrient cycling within forests is critical for long term rdaintenatol bos produlgo ty

and stability. It involves nutrient uptake, utilization and accumulation by vegetation,
together with the return of nutrients to the soil through litterfall, leaching and root
sloughing. Comparisons of nutrient cycles involve considerations HK Spe iesrelable ss.
to obtain nutrients from soils with low nutrient status and then rétain-the-nutrients.___
within systems (Turner, 1975). In order to assess turnover of nutrients, various indices
have been developed. For example, turnover of litter on forest floor has been compared

by using a ‘k’ factor which relates the input of litter (L) to the mass of litter (F) on the
forest floor. The ‘k’ factor = L/F and assumes a steady state forest floor weight (Jenny e¢

al., 1949; Olson, 1963; Richards and Charley, 1977) and gives an indication of the rate of

loss (decomposition) by the litterfall in relation to accumulation on the forest floor.

Relative efficiency of nutrient acquisition from soil, nutrient utilization requirements
and efficiency of redistribution of nutrients can be assessed within the tree component of
an ecosystem. These comparisons are difficult, but some can be made. For example, an
index of nutrient utilization within a tree is often obtained using foliage nutrient con-
centrations (Lambert and Turner, 1983; Lambert et a/., 1983). Nutrient redistribution
may be estimated as withdrawal of nutrients, both during leaf abscission (Ashton, 1976;
Attiwill, 1980; Turner and Lambert, 1983) and in heartwood formation (Lambert,
1981). Such estimates have been made in only a few forests in Australia and have been
carried out predominantly in stands dominated by a single species (Hingston et al., 1979;
Attiwill, 1980; Turner and Lambert, 1983) where monthly leaf litterfall data were com-
pared with those for live leaf material on the trees. In studies of forest stands including a
variety of species, and particularly in conditions where organic matter decomposition
and tissue leaching can be quite rapid, the use of monthly litterfall samples becomes
inappropriate.

Subtropical rainforests are associated with relatively fertile soils (Baur, 1957; Webb,
1969; Lambert et a/., 1983) whereas eucalypt forests are on soils with much lower fertility
(Baur, 1957; Webb, 1969; Turner and Kelly, 1981). Subtropical rainforests (Baur, 1965)
have high species diversity with often in excess of 30 species ha" in the overstorey. They
are notable in northern N.S.W. for the absence of Eucalyptus species, a genus which

PROG. LINN. SOC. N.S.W., 111 (1), 1989

2 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES

dominates most other coastal and tableland forest types in N.S.W. During a programme
of study in a subtropical rainforest located on the New South Wales Border Ranges, leaf
material was sampled from a range of species 1n order to obtain indices of nutrient distri-
bution and cycling patterns within this forest. Fresh litter was specifically sampled to
provide estimates of nutrient redistribution in various species. These data were com-
bined with differences in heartwood and sapwood nutrient concentrations and
compared with similar data from coastal sclerophyll forests.

STUDY SITE

The study site was originally described by Lambert e¢ al. (1983) and was located in
forests of the Border Ranges (153°E, 28°38’S), west of Murwillumbah. The altitudinal
range of the forest is 600-1200m above sea level. Annual rainfall is 3000mm. Plots were
selected from within subtropical rainforest growth experiments (Burgess et al., 1975;
Horne and Gwalter, 1982). Soils from the region are derived from Tertiary volcanic
rocks of the Mt Warning Shield and are predominantly of basaltic composition
(Stevens, 1976). The basalts have given rise to kraznozems, that is, deep well-structured
red clay loams with clay sub-soils having a relatively uniform appearance and depth
(Beckman and Thompson, 1976). The soils are high in nutrients, particularly when
compared with soils supporting sclerophyllous species (Lambert et al., 1983).

METHODS

Trees were sampled for foliage and wood during logging operations near the study
site. Within a species, nutrient variability in foliage concentration was found to be low
(Lambert et al/., 1983; Lambert and Turner, 1986). Foliage samples were bulked from
within the crown but only fully-formed leaves were used; that is, very young or damaged
leaves were omitted. Samples were placed in paper bags, oven dried at 70°C, ground
and analysed for various chemical elements (Lambert, 1983). The results for overstorey
and understorey trees have been reported elsewhere. Wood discs were taken from the
stem approximately 1m above the ground. These were air dried and then separated into
bark, sapwood and, where present, heartwood. The individual components were
ground and analysed for the same chemical elements as the foliage samples (Lambert,
1983).

Freshly-fallen leaf samples were sampled beneath the crown of selected species
within the research plot and were those recently fallen so that there was minimal effect
due to decomposition and leaching. Where leaves were caught on understorey vegeta-
tion, they were preferentially selected so that ground contact was minimized. Acquiring
sufficient suitable material of many species was difficult and hence single, bulked
samples were used rather than replicates. The leaf litterfall samples were dried, ground
and analysed in the same way as foliage samples.

A list of the common names, scientific names and authorities of species sampled on
the study site is given in the Appendix.

RESULTS

Results of the analyses are presented in detail in Table 1; trends are summarized in
Table 2. Changes in concentration of nitrogen and phosphorus between overstorey leaf
and litterfall were relatively minor in most cases. There were obvious exceptions, such as
Dendrocnide excelsa, Solanum aviculare and Solanum mauritianum, these tending to be very
high in nutrients initially. The other exception was Orites excelsa which has relatively low
requirements for both phosphorus and nitrogen and appears to be also efficient at
retranslocation, that is, there is high nutrient removal by retranslocation even in a situ-

PROC. LINN. SOC. N.SW., 111 (1), 1989

M. J. LAMBERT AND J. TURNER

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PROC. LINN. SOC. N.S.W., 111 (1), 1989

NUTRIENT REDISTRIBUTION IN RAINFOREST TREES

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PROC. LINN. SOC. N.S.W., 111 (1), 1989

M. J. LAMBERT AND J. TURNER 9)

ation of high nutrient availability (Kelly e¢ a/., 1983). Calcium typically increased in con-
centration in the litter of most species, an exception being Orites excelsa. The patterns of
other elements were much more irregular (Table 2). Potassium and manganese tended
to decline and accumulate respectively. Caldcluvia paniculosa and Orites excelsa accumu-
lated aluminium in the foliage but the concentration of this element was lower in the
litter, the Orites excelsa to a higher degree possibly showing different patterns of handling
an antagonistic element. Of the 48 samples assessed, heartwood was detectable in only
12 and in these instances relatively few had significant retranslocation of nutrients
(Lambert et al., 1983).

Within the non-woody understorey species, nutrients were retranslocated between
young and old tissues as in the woody species (Table 3). Cyathea australis, for example,
redistributed 45% of phosphorus in the aging of fronds. Archontophoenix cunninghamiana
was an efficient retranslocater of phosphorus (65%) and potassium (96%), although
there may be a higher proportion leached as it is difficult to estimate when foliage of this
species can be classified as litterfall as the older foliage hangs down next to the stem.
Asplenium nidus retranslocated much smaller quantities during senescence.

DISCUSSION

Estimates of nutrient retranslocation are relative measures and to understand the
ecological significance of nutritional patterns, comparisons with other forest types have
been used. Within the subtropical rainforest, retranslocation of nutrients within foliage
prior to abscission appears to be relatively low for most overstorey species, however,
exceptions were: Dendrocnide excelsa, this species having fairly high nutrient require-
ments; Orites excelsa, which is an aluminium-accumulating species and appears to have
an efficient system of retranslocation; and two solanaceous species (Solanum aviculare and
Solanum mauritianum which invade and grow immediately after soil disturbance and live
for only a relatively short time. It appears they have both high nutrient demands and
efficient retranslocation. The level of retranslocation may be compared with that in
sclerophyllous species and coachwood which generally grow on nutritionally poorer soils
than the subtropical rainforest (Table 4). In these species, phosphorus, nitrogen, mag-
nesium and potassium are all retranslocated, while calcium and aluminium are
accumulated. The calcium pattern appears typical of mature foliage in many trees.
Further, there were consistently high removals of nutrients in these species during heart-
wood development. The pattern of heartwood development and retranslocation was
either absent or very low in the majority of species sampled in this rainforest.

Generally, nutrient redistribution is an important component, along with uptake,
in fulfilling forest stand nutrient requirements (Turner and Lambert, 1983). However, if
the general principle is that the subtropical rainforest tree species generally redistribute
nutrients at a low level, whereas the sclerophyllous and cool temperate rainforest species
are more efficient at redistribution, is this a function of generally higher soil nutrient
availability? Further, iflarge fertilizer quantities were applied to eucalypt species or they
were located on more fertile soils, would heartwood production be reduced or litter
retranslocation lowered? Eucalypts growing naturally across a range of fertilities do not
have significantly different foliage nutrient concentrations (Lambert and Turner, 1983)
or nutrient redistribution patterns. This suggests that soil nutrients are not controlling
the patterns but rather that they are evolutionarily determined. The reverse situation of
growing subtropical rainforest trees on poor sites is harder to test, mainly because it
appears that soil nutrients are delineators of species (Turner and Kelly, 1981) and hence
rainforest species are out-competed on poor sites.

Within the subtropical rainforest there were some species such as Orites excelsa

PROG. LINN. SOG. N.S.W., 111 (1), 1989

NUTRIENT REDISTRIBUTION IN RAINFOREST TREES

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PROC. LINN. SOC. N.S.W., 111 (1), 1989

8 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES

TABLE 4

Published information for various Australian forest species on redistribution from foliage during abscission

Forest Reference

Type

IP Ca Meg K Al
(ppm)

Wet Sclerophyll
E. grandis — foliage 895 4800 2690 6450 80 Turner and
— litter 340 7065 2160 1955 230 Lambert
— change -0.61 =599 +2265 -530 -4495 +150 | (1983)
E. regnans* — fohage 0.79) 780 5730 5730 Ashton
— litter 0.87 300 5450 1240 (1976)
— change (0), 112 -480 -280 -4490)
E. pilularis — foliage 1.21 620 3420 3465 3810 120 | Turner and
— litter 070 300 WAI ASSO. lleXO) 330 | Kelly
— change -0.51 -320 +9190 -1075 -2670 +210 | (1981)
Lophostemon — foliage 1.61 1205 V525) 2735 11035 195 Turner and
confertus — litter 0.81 529) 12220 2580 1750 820 | Kelly
— change -0).80 -680 +2695 159 -9285 +625 | (1981)
Rainforest
Ceratopetalum — foliage 1.10 505 10365 2990 4540 TOUS Turner and
apetalum = litter 0.71 210 7890 1740 1030 8200 | Kelly
— change =), 39 -295 -27495 -1250 -3510 +925 (1981)
Nothofagus — foliage 1010 Ashton
cunningham — litter _ 300 (1976)
— change -710
Athosporia — foliage 1500 Ashton
moschatum — litter 890 (1976)
— change -630
Dry Sclerophyll
E. sveberi — foliage 440 Ashton
— litter 110 (1976)
— change -330

4 Qnd year leaf and leaf litter analyses for a mature stand.

which gave a similar pattern to the sclerophyllous pattern of redistribution. Whether
this is an indication that the species had evolved on poorer soils and now survive within
the subtropical rainforest is difficult to ascertain, but it is suggested that the com-
bination of high litter and heartwood redistribution efficiencies, particularly for phos-
phorus, is indicative of evolution on soils with low nutrient status.

A further question arises as to the relationship between redistribution during leaf
abscission and that during heartwood formation for the rainforest species. For most
nutrients there was no relationship, but for phosphorus and nitrogen, there was an
inverse relationship (Fig. 1). That is, as the proportion of phosphorus redistributed in
leaves increased, the proportion redistributed in heartwood formation or its formation
at all, decreased. The exception to this was Orites excelsa which, as noted above, tends to
have unusual patterns of nutrient utilization compared with other species.

The general pattern for phosphorus in rainforest trees, is that as phosphorus
becomes more efficient at redistribution in one type of tissue, it becomes less efficient in
another. The species which most noticeably form heartwood are not redistributing
during leaf senescence. This is the opposite pattern to that found in Eucalyptus and other
sclerophyllous species (Fig. 1) where there is simultaneous redistribution from leaf litter

PROG. LINN. SOC. N.S.W., 111 (1), 1989

M. J. LAMBERT AND J. TURNER

Ko)

ROSE RUS" fe aa
80
60
40
20

-40 -20 0 20 40 60
LEAF LITTER REDISTRIBUTION (%)

Fig. 1. Relationship between redistribution of phosphorus during heartwood formation and leaf abscission
(¢ — rainforest species; x — eucalyptus and sclerophyllous species; L] — Ceratopetalum apetalum; O — Orites
excelsa).

and heartwood. Orites excelsa relates closely to the ‘sclerophyllous’ pattern which includes
E. pilularis, E. dives, E. maculosa, E. rossu, E. rubida, E. obliqua, E. grandis, Lophostemon
confertus and Casuarina torulosa, these being the species where mature green leaf, leaf
litterfall, sapwood and heartwood concentrations were available. The phosphorus re-
distribution pattern in coachwood, however, was similar to that found generally in the
subtropical rainforest species even though this species grows in differently structured
rainforest (Baur, 1965).

Comparisons with other species (Table 4) were based on results for mature leaves
on the tree and fresh litter using comparable sampling techniques. Fully-developed
younger leaves have nutrient concentrations different to those in older leaves, so that
there are different patterns of redistribution taking place within the crown. Further,
there are different patterns between different-aged forests of the same species (Ashton,
1975). The pattern for nitrogen was similar to that for phosphorus for the rainforest
species. Orites excelsa was again found to form a different pattern. Nitrogen was generally
not as efficiently redistributed as phosphorus during heartwood formation. Ceratopetalum
apetalum and Lophostemon confertus in this case followed the rainforest pattern, while the
eucalypts were differently distributed.

While for most nutrients there are relationships between foliage litter and heart-
wood redistribution which may be described as specific to species/site, phosphorus and
nitrogen have given separate patterns of nutrient cycling. In rainforests, where phos-
phorus and nitrogen are usually readily available, a certain amount of ‘energy’ is
apparently expended in retaining nutrients in biomass and hence not all tissues are
affected. In the case of the lower phosphorus sclerophyllous forest, all available phos-
phorus is apparently redistributed, this being the primary limiting nutrient. The
pattern of Orites excelsa possibly indicated that it evolved in a low phosphorus
environment.

References
ASHTON, D. H., 1976. — Phosphorus in forest ecosystems at Beenak, Victoria. /. Ecol. 64: 171-186.
ATTIWILL, P. M., 1980. — Nutrient cycling in a Eucalyptus obliqua (L-Hérit.) forest — IV. Nutrient uptake and
nutrient return. Aust. J. Bot. 28: 199-222.

PROG. LINN. SOC. N.S.W., 111 (1),

10 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES

Baur, G. W., 1957. — Nature and distribution of rain-forests in New South Wales. Aust. J. Bot. 51: 190-222.

— , 1965. — Forest types in New South Wales. For. Comm. N.S.W. Res. Note No. 17.

BECKMAN, G. G., and THOMPSON, C. H., 1976. — The soils. In The Border Ranges — a land use conflict in regional
perspective (cds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Qucensland.

BurGess, I. P., FLoyp, A., KikKAwA, J., and PATTIMORE, V., 1975. — Recent developments in the silvi-
culture and management of subtropical rainforest in N.S.W. Proc. Ecol. Soc. Aust. 9: 74-84.

HINGSTON, F. J., TURTON, A. G., and DIMMOCK, G. M., 1979. — Nutrient distribution in Karri (Eucalyptus
diversicolor F. Muell.) ecosystems in southwest Western Australia. For. Ecol. Managem. 2: 133-158.
HORNE, R., and GWALTER, J., 1982. — The recovery of rainforest overstorcy following logging. I. Subtropi-

cal rainforest. Aust. For. Res. 13: 29-44.
JENNY, H., GesseL, S. P., and BINGHAM, F. T., 1949. — Comparative study of decomposition rates of organic
matter in temperate and tropical regions. Soz/ Scz. 68: 419-432.
KELLY, J., LAMBERT, M. J., and TURNER, J., 1983. — Available phosphorus forms in forest soils and thcir
possible ecological significance. Commun. Soil Sci. Plant Anal. 14: 1217-1234.
LAMBERT, M. J., 1981. — Inorganic constituents in wood and bark of New South Wales forest tree species.
For. Comm. N.S.W., Res. Note 45. 43 pp.
—., 1983. — Methods for chemical analysis. For. Comm. N.S.W. Tech. Pap. 25. Third Edition. 187 pp.
, and TURNER, J., 1983. — Soil nutrient-vegetation relationships in the Eden area, N.S.W. III. Foliage
nutrient relationships with particular reference to Eucalyptus sub genera. Aust. For. 46: 200-209.
, and , 1986. — Nutrient concentrations in foliage of species within a New South Wales sub-tropical
rainforest. Ann. Bot. 58: 465-478. .
, and KELLY, J., 1983. — Nutrient relationships of tree species in a New South Wales sub-tropical
rainforest. Aust. For. Res. 13: 91-102.
OLSON, J. S., 1963. — Energy storage and the balance of producers and decomposers in ecological systems.
Ecology 44: 322-331.

RICHARDS, B. N., and CHARLEY, J., 1977. — Carbon and nitrogen flux through native forest floors. In
Nutrient cycling in indigenous forest ecosystems: 65-81. Perth: C.S.I.R.O. Division of Land Research and
Management.

STEVENS, N. C., 1976. — Geology and landforms. In The Border Ranges — a land use conflict in regional perspective
(eds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Queensland.
TURNER, J., 1975. — Nutrient cycling in a Douglas-fir ecosystem with respect to age and nutrient status.
Seattle, Washington: University of Washington, Ph.D. thesis, unpubl.
, and KELLY, J., 1981. Relationships between soil nutrients and vegetation in a north coast forest, New
South Wales. Aust. For. Res. 11: 201-208.
, and LAMBERT, M. J., 1983. — Nutrient cycling within a 27-year-old Eucalyptus grandis plantation in
New South Wales. For. Ecol. Managem. 6: 155-168.
WEBB, L. J., 1969. — Edaphic differentiation of some forest types in eastern Australia. II. Soil chemical
factors. J. Ecol. 57: 817-830.

APPENDIX

List of common names, scientific names and authorities of species from the study site

Common name

Bangalow palm
Birds nest fern

Botanical name

Archontophoenix cunninghamiana (H. Wendl.) H. Wend. et Drude
Asplenium nidus L.

Black booyong Heritiera actinophylla (F. M. Bail.) Kosterm.
Bollygum Neolttsea reticulata (Meisn.) F. Muell.
Brushbox Lophostemon confertus (R. Br.) Peter G. Wilson e J. T. Waterhouse
Coachwood Ceratopetalum apetalum D. Don

Corkwood Caldcluvia paniculosa (F. Muell.) Hoogl.
Doughwood Euodtia micrococca F. Muell.

Giant stinging tree Dendrocnide excelsa (Wedd.) Chew
Kangaroo apple Solanum aviculare Forst. f.

Lace fern Nephrolepis sp.

Pigeonberry ash Cryptocarya erythroxylon Maiden et Betche
Prickly ash Orites excelsa R. Br.

Red carabeen Geissois benthamiana F. Muell.

Rosewood Dysoxylum fraseranum (A. Juss.) Benth.
Teak Flindersia australis R. Br.

Tree fern Cyathea australis (R. Br.) Domin

White booyong Heritiera trifoliolata (F. Muell.) Kosterm.

Wild tobacco tree
Yellow carabeen

Solanum mauritianum Scop.
Sloanea woollsii F. Muell.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

New Occurrences of Culmacanthid
Acanthodians (Pisces, Devonian) from
Antarctica and southeastern Australia

G. C. YOUNG

YOUNG, G. C. New occurrences of culmacanthid acanthodians (Pisces, Devonian)
from Antarctica and southeastern Australia Proc. Linn. Soc. N.S.W. 111 (1), 1989:
11-24.

Two new spccies of the acanthodian fish Culmacanthus Long 1983 are described
from the lower part of the Aztec Siltstone of southern Victoria Land. Antarctica, and
Facies 3 of the Boyd Volcanic Complex of southeastern New South Wales. Both are
represented only by cheek plates. That of C. antarctica sp. nov. has distinctive ornament,
a longer lateral than dorsal lamina, and the infraorbital sensory groove passing off the
ventral margin of the plate. C. pambulensis sp. nov. resembles the type species C. stewarti
Long in the shape of the posterior margin of the check plate and the ventral course of the
infraorbital sensory canal, but differs in its proportions, the shape of the anterior
margin, and the much finer dermal ornament. C. antarctica sp. nov. is considered to be
the oldest (?late Middle Devonian) because it is associated with turiniid thelodont
scales. The two other species of Culmacanthus occur with bothriolepid and phyllolepid
placoderms in assemblages considered to be early Late Devonian (Frasnian). The
specific differences described herein suggest however that they are not precise cor-
relatives. Culmacanthid acanthodians are only known from southeastern Australia and
southern Victoria Land, a distribution pattern previously noted in chondrichthyans
and placoderms from the same faunas, and suggesting that Culmacanthus was an cast
Gondwanan endemic.

G. C. Young, Division of Continental Geology, Bureau of Mineral Resources, PO. Box 378,
Canberra, Australia 2601; manuscript received 15 December 1987, accepted for publication
24 August 1988.

INTRODUCTION

The acanthodians are a major group of Palaeozoic gnathostome fishes, with a fossil
record from Silurian to Permian. During the Devonian Period they were widely dis-
tributed in both non-marine and marine aquatic environments, and their tiny scales
and characteristic fin-spines are commonly represented in microvertebrate assem-
blages. However articulated specimens are much less common, and knowledge of
acanthodian morphology is based mainly on a few localities in Europe and North
America which have yielded well-preserved complete specimens (for a comprehensive
review of the group see Denison, 1979).

A. S. Woodward was the first to describe Australian acanthodians, from the Lower
Carboniferous Mansfield group of Victoria (Woodward, 1906), and he also published
the first descriptions of the group from Antarctica (Woodward, 1921). Isolated acantho-
dian scales and spines occur widely in Devonian rocks of eastern and central Australia
(Fig. 1), and in recent years some well-preserved articulated acanthodians have been
described from southeastern Australia (e.g. Long, 1983a, 1986a). The best-known
locality is Mount Howitt in east central Victoria, which is the type locality for the genus
Culmacanthus Long 1983, of which additional material is described in this paper. With
these two new species the genus Culmacanthus is now known from three localities in
southeastern Australia, and one locality in Antarctica (Fig. 1). Its distribution pattern
matches that seen in several other taxa of Devonian fishes, the palaeogeographic sig-
nificance of which is discussed below. All described specimens are housed in the Com-
monwealth Palaeontological Collection (prefix CPC), Bureau of Mineral Resources,

PROC. LINN. SOC. N.S.W., 111 (1), 1989

12 DEVONIAN FOSSIL FISHES

abs Als
MR
WC 4 A ARS
AUSTRALIA

EAST ANTARCTICA

20/09/172

Fig. 1. Reconstruction of East Antarctica against Australia, modified from the Gondwana reconstruction of
Lawyer and Scotese (1987). showing the main localities of Devonian acanthodian fishes in the southwest
Pacific region. Numbered localities are the only known occurrences of culmacanthid acanthodians, as dealt
with in this paper; 1, Mount Crean, Lashly Range, southern Victoria Land (Culmacanthus antarctica sp. nov.);
2, Mount Howitt, cast central Victoria (Culmacanthus stewart: Long); 3, Freestone Creek, east Gippsland,
(culmacanthid indet.); 4, Pambula River, southeastern New South Wales (Culmacanthus pambulensis sp. nov.).
Other abbreviations are: B, Broken River, Queensland; BD, Burrinjuck Dam, New South Wales; C, Cobar
area, Darling Basin; DS, Dulcie Syncline, Georgina Basin, EM, Crashsite Quartzite, Ellsworth Mountains;
G, Grampians, western Victoria; H, Horlick Formation, Ohio Range; M, Munyarai, Officer Basin; MBL,
Marie Byrd Land, West Antarctica, MR, Macdonnell Range, Amadeus Basin; NNZ, New Zealand, North
Island; R, Reefton, New Zealand, South Island; RS, Ross River Syncline, Amadeus Basin; T, Tasmania; TS,
Toko Syncline, Georgina Basin; WC, Wilson Cliffs, Canning Basin.

Geology and Geophysics, Canberra, A.C‘T. Material of the type species Culmacanthus
stewarti Long is housed in the Museum of Victoria (prefix NMP).

PREVIOUS INVESTIGATIONS

The first samples of the Antarctic Aztec fish fauna were collected by F. Debenham
on the British Antarctic “Ierra Nova expedition of 1910-13, from moraine material at

PROC. LINN. SOG. N.S.W., 111 (1), 1989

G. C. YOUNG 13

Granite Harbour on the coast of southern Victoria Land. They were described by
Woodward (1921), who identified eight taxa of Devonian fishes, including isolated
acanthodian scales which were said to resemble those of Chezracanthus murchisoni from the
Middle Devonian of Scotland. /n situ remains of the Aztec fauna were not discovered
until the Trans-Antarctic Expedition of 1955-58 (see Gunn and Warren, 1962), when
Devonian fish remains were collected from the Aztec Siltstone at three localities (Lashly
Mountains, Mount Feather, Boomerang Range) in the region of the Skelton Névé of
southern Victoria Land (locality 1, Fig. 1). Other fish remains from the same general
area were recorded by Matz and Hayes (1966) and Matz, Pinet and Hayes (1972; their
‘Fortress Formation’) from west Beacon Heights, Mount Fleming, Mount Crean, and
Aztec Mountain, and by Helby and McElroy (1969) from Aztec Mountain. Gunn and
Warren's collection was described by White (1968). The most extensive collections of
Devonian fishes from the Aztec Siltstone were made by Antarctic expeditions from Vic-
toria University of Wellington in 1968-69 (see McKelvey et al., 1972), and in 1970-71
(VUWAE 15), when the specimen described below was collected. Detailed measured
sections through the Aztec Siltstone at most of the fossil localities are given in Askin et al.
(1971) and Barrett and Webb (1973). Recent publications dealing with or discussing the
Aztec fish fauna include Ritchie (1971a, b, 1974, 1975). Young (1982, 1988), and Grande
and Eastman (1986). A detailed locality map and a summary of the 24 known localities
for the Aztec Siltstone fish fauna are given in Young (1988).

In the material collected by Gunn and Warren, White (1968) identified several
types of acanthodian spines. Ribbed spines from the upper fossiliferous horizon in the
Boomerang Range were referred to a new species (G. warren) of the genus Gyracanthides,
originally described by Woodward (1906) from the Early Carboniferous of Mansfield in
Victoria. A new genus and species (Antarctonchus glacialis White) was erected for long
slender acanthodian spines with numerous strong longitudinal ribs. White also deter-
mined as an acanthodian some small spine fragments originally described by Woodward
(1921) as an antiarch (Byssacanthoides debenhami Woodward). In addition to these named
taxa from the Middle-Late Devonian, V. T. Young (1986) has referred to the genus
Machaeracanthus two types of Early Devonian acanthodian spines from the Horlick
Formation in the Ohio Range of the southern Transantarctic Mountains, and a similar
specimen is recorded from the Ellsworth Mountains (H, EM, Fig. 1).

In southeastern Australia acanthodian remains have been known from Victoria for
some time (Woodward, 1906; Hills, 1931). More recently Long (1983a) described
Culmacanthus stewart: from the Mount Howitt fish locality, and figured one incomplete
specimen from Freestone Creek in eastern Victoria as a probable culmacanthid (locali-
ties 2, 3, Fig. 1). A second acanthodian from the Mount Howitt fauna (Howzttacanthus)
was described by Long (1986a). A summary of the Victorian Devonian fish localities and
their biostratigraphy is given in Long (1983b: fig. 1). Placoderms associated with
Culmacanthus are also described by Long (1984) and Long and Werdelin (1986).

Over the state border, in southeastern New South Wales, Devonian fish remains
were discovered in the Eden-Pambula area in 1978 (locality 4, Fig. 1), and a preliminary
report (Young, in Fergusson et al., 1979) identified four assemblages ranging in age
probably from iatest Middle Devonian (Givetian) through to the end of the Late
Devonian (Famennian). One specimen, first described as a possible osteichthyan plate
(Fergusson et al., 1979: 103), is referred below to Culmacanthus on the basis of Long’s
(1983a) description of this genus. Further collecting in 1979 produced the new speci-
mens described below. Other described elements of the Devonian fish faunas from the
south coast of New South Wales include sharks from the Bunga Beds near Bermagui
(Young, 1982), and an asterolepid antiarch from the Pambula River area (Young, 1983).
Elements still to be described include bothriolepid and remigolepid antiarchs, phyllo-

PROG. LINN. SOC. N.S.W., 111 (1), 1989

14 DEVONIAN FOSSIL FISHES

lepid placoderms, and rhipidistian and other acanthodian remains. A locality map of
the region is given in Young (1983: fig. 1).

SYSTEMATIC DESCRIPTIONS
Subclass ACANTHODII Owen
Order CLIMATIIDA Berg
Suborder DIPLACANTHOIDEI Miles 1966
Family CULMACANTHIDAE Long 1983
Genus Culmacanthus Long

Remarks: The above taxa are defined in Denison (1979) and Long (1983a).

Culmacanthus antarctica sp. nov.
(Figs. 2A, 3A)

1986a. ‘culmacanthid cheek plates’ (pars) Long, p. 13.
Holotype: CPC 26579, a right cheek plate preserved in part and counterpart.
Locality: Mount Crean, east side of the Lashly Range (77° 53'S, 159° 33’E), southern
Victoria Land, Antarctica (locality 8 of Young, 1988: fig. 3).
Horizon: From the lower part of the Aztec Siltstone, within 40 m of the base of the for-
mation (unit 5 of section L2 measured by Askin et al., 1971; for a detailed discussion of
the stratigraphy of the Mount Crean section see Young, 1988).
Diagnosis: A Culmacanthus possessing a cheek plate in which the infraorbital sensory
canal passes off the ventral margin, the anteroventral corner is pronounced, the ventral
lamina is longer and deeper than the dorsal lamina, and the plate is fairly flat,
ornamented with closely spaced ridges and elongate tubercles with a flat external
surface, and has a breadth/length index of about 79.
Remarks: Long (1983a) did not present a diagnosis for the type species C. stewart, but
compared to the Antarctic specimen there are obvious differences in the form of the
cheek plate. In the type species the coarser ornament of more widely spaced ridges have
a rounded rather than flat dorsal surface, the anterior margin of the plate is distinctly
notched (Fig. 3B), there is a more pronounced angle between the dorsal and lateral
laminae, the dorsal lamina is longer than the lateral, and the latter has a concave exter-
nal surface. The most obvious difference is in the course of the main sensory groove,
with both dorsal and ventral sections passing off the anterior margin of the plate.

The articulated remains of C. stewart: show that Culmacanthus was a diplacanthoid
acanthodian with two dorsal fin-spines, paired pectoral and pelvic spines, and an anal
spine (Fig. 5). These spines are ornamented with approximately nine coarse longitudi-
nal ribs, and they have a deeply inserted base which is finely striated (Long, 1983a). It is
possible therefore that some of the similar acanthodian spines from the Aztec Siltstone
named by White (1968) as Antarctonchus may also belong to Culmacanthus. Long (1983a)
has already suggested this for the isolated spines from Freestone Creek in Victoria
named Striacanthus by Hills (1931). However, in the absence of articulated material
demonstrating this, and because the histology of Culmacanthus spines is at present
unknown, I follow Long’s (1983a) procedure here, and leave Antarctonchus as a form
genus for isolated spines, until such time as new material or more detailed study can
demonstrate that the spines and dermal cheek plates come from the same fish.
Description: The holotype and only referred specimen 1s an isolated dermal cheek plate
preserved in part and counterpart. The external impression was cleaned in dilute
hydrochloric acid to remove the bone, and the distinctive ornament is well displayed in a
latex cast (Fig. 2A). The counterpart is preserved as bone except for the posterolateral
corner, which is an impression of the visceral surface. All margins of the plate are

PROC. LINN. SOG. N.S.W., 111 (1), 1989

G. C. YOUNG 15

Fig. 2. Right cheek plates of Culmacanthus in external view (latex casts whitened with ammonium chloride). A,
C. antarctica sp. nov., holotype, CPC 26579 (x4). B-D, C. pambulensis sp. nov.; B, holotype, CPC 26580 (x4); C,
CPC 26581 (x3); D, CPC 26582 (x4).

complete. The orientation of such isolated cheek plates is not readily determined, but
following the work of Long (1983a) on the articulated Mount Howitt material of C.
stewart, the position of the short sensory groove (ifc.b, Fig. 3A) shows this to be a right
dermal cheek plate. It is about 21.5mm long, and relatively deep (breadth/length index
of 79). The anterior margin lacks the distinct orbital notches seen in the type species
(Fig. 3B), and slopes anteroventrally to a pronounced anteroventral corner, behind
which the infraorbital sensory groove passes off the ventral margin of the plate.

The plate has a slight flexure at the level of the posterior angle of the infraorbital
sensory groove (ifc, Fig. 3), as previously described by Long (1983a), but overall it is a
much flatter plate than in the Victorian species (based on NMP 159838). The inflexion
of the infraorbital groove is placed high on the plate, which gives it a much larger lateral
than dorsal lamina. The short posterodorsal sensory groove (ifc.b), as described for C.
stewarti, was interpreted by Long (1983a) as possibly part of the preopercular sensory
groove, but this groove in fishes normally passes posterolaterally across the cheek
towards the mandibular joint. It is considered more likely that the short groove on the

PROG. LINN. SOG. N.S.W., 111 (1), 1989

16 DEVONIAN FOSSIL FISHES

cheek plate of Culmacanthus is equivalent to the ‘postero-median branch of the infra-
orbital canal’ of Diplacanthus identified by Watson (1937; ‘central sensory line’ of Miles,
1966).

Fig. 3. Cheek plates of Culmacanthus restored in external view. A, a right plate of C. antarctica sp. nov., after the
holotype, CPC 26579; B, C. stewart: Long, a left plate, restored after a latex cast of NMV P159838 (see Long
1983a: fig. 4B); C, aright plate of C. pambulensis sp. nov., after the holotype, CPC 26580. an, anterior notch;
dep.p, posterior depression; ifc, infraorbital sensory groove; ifc.b, posterodorsal (‘central’) branch of infra-
orbital sensory groove; pn, posterior notch.

The distinctive ornament of C. antarctica sp. nov. comprises cusp-like tubercles
which change into elongate ridges towards the posterior margin. In front of the sensory
groove the surface of each cusp slopes upwards from the bone surface in a posterodorsal
direction, at which a point may be developed, to give some cusps a triangular shape.
Behind the sensory groove the more elongate cusps and short ridges have their pointed
ends directed posteriorly, and are separated by deep, narrow grooves. Most of the ridges
and tubercles are ornamented with up to four very fine ridges which may branch
towards the anterior. Typically the elongate primary ridges of the ornament which are
most pronounced on the posterior half of the plate have a flat or slightly concave upper
surface, with the edges clearly marked by two fine ridges. Compared to C. stewartt (NMP
159838, Fig. 3B) the ornament is more finely sculptured, with the ridges and tubercles
more closely spaced. In the type species (Long, 1983a, fig. 4A, B) the ornament is

PROC. LINN. SOG. N.S.W,, 111 (1), 1989

G. C. YOUNG Vi)

coarser, with wider intervening spaces, and scattered tubercles along the ventral
margin. However some fine striations are also visible on the more elongate tubercles and
ridges, and in larger specimens there are fewer tubercles and longer ridges (Long,

1983a, fig. 2B).

Culmacanthus pambulensis sp. nov.
(Figs. 2B-D, 3C, 4)
1979. ‘osteichthyan, with an ornament of delicate subparallel ridges’ Young, in Fergus-
son et al., p. 103.
1983a. ‘dermal cheek plate. . . from Pambula River’ Long, p. 54.
1986a. ‘culmacanthid cheek plates’ (pars) Long, p. 13.

Fig. 4. Culmacanthus pambulensis sp. nov., cheek plates in external view. A, a right plate, after CPC 26581; B, a
left plate, after CPC 26583. For abbreviations see caption to Fig. 3.

PROC. LINN. SOG. N.S.W., 111 (1), 1989

18 DEVONIAN FOSSIL FISHES

Name: Abbreviated from the nearby town of Pambula, N.S.W.

Holotype: CPC 26580, a right cheek plate.

Other Material: Right (CPC 26581, 26582) and left (CPC 26583) cheek plates, and
another ornamented fragment (CPC 17004).

Locality: Grid Reference 7510E 59081N on the Pambula 1:25000 Sheet, about 5km west
of the town of Pambula on the south coast of New South Wales (see Young, 1983: fig. 1).
Horizon: Facies 3 of the Boyd Volcanic Complex, as described by Fergusson et al. (1979).
Diagnosis: A Culmacanthus possessing a fairly flat cheek plate in which the infraorbital
sensory canal passes off the anterior margin, the anterior margin has only shallow or in-
distinct notches, the ventral lamina 1s considerably deeper than the dorsal lamina, the
closely spaced ornament comprises fine ridges and elongate tubercles, and the
breadth/length index varies between at least 68 and 94.

Remarks: Apart from the obvious and consistent differences in ornament, the three
measurable specimens of C. pambulensis are broader than in the type species, although
for the smallest example described below (CPC 26580) this difference is only slight.
Long (1983a: 56) regarded such differences in proportion to be generic rather than
specific features, but the variability in proportions is clear in this new species, and the
other obvious similarities indicate that these specimens are properly referred to the
genus. C. pambulensis resembles C. antarctica rather than C. stewarti in the flatter cheek
plate, the longer and steeper disposition of the ventral section of the infraorbital sensory
groove, and the absence or slight development of notches on the anterior margin. It
differs from C. antarctica but resembles C. stewarti in the fact that the infraorbital groove
passes off the anterior margin of the plate. Long (1983a) noted other specimens from
Freestone Creek in eastern Victoria with broader proportions than the type species, as
in C. pambulensis sp. nov., but that material is at present too poorly known to permit
useful comparisons.

Description: All specimens are preserved as external moulds, which have been studied
by latex casting. The holotype (Figs 2B, 3C) is the smallest specimen (length 14mm),
and the most complete. The infraorbital groove has an attenuated ventral section, not
seen in other examples, and interpreted as an individual variation. The posterodorsal
branch is a short separate groove (ifc.b, Fig. 3C) as in other examples with this part
preserved. Compared to C. stewart: (represented by NMP 159838) the cheek plate is
rather flat. The lateral lamina is slightly concave near the posterior corner (dep.p), and
there may be another shallow depression running anteroventrally beneath the ventral
section of the infraorbital canal, but otherwise both laminae of the plate are gently con-
vex. The posterior depression is seen in all available specimens of this species (Fig. 2B-
D). Two larger specimens are unusually broad (Fig. 4), although their posterior margins
are somewhat irregular. This may be due to incomplete preservation, but an irregular
margin abutting against small plates at the posterior margin of the gill cover is con-
sidered more likely. It is possible that breadth of the plate increased allometrically with
size, but evidence against this is provided by C. stewartz, where the largest plate (Long,
1983a: fig. 2) is also the most elongate (erroneously stated as ‘twice as broad as long’ by
Long, 1983a: 56). In both CPC 26581 and 26583 the lateral lamina projects posteriorly,
as in C\ antarctica described above, and all specimens with the region preserved show a
shallow notch on the posterodorsal margin (pn, Figs 3C, 4). The general similarity of the
delicate ornament in the two largest examples (Figs 2C, 4B) to other specimens from this
locality suggests that they are conspecific, and that proportions of the cheek plate are
more variable than previously thought. In the type species only three cheek plates have
been described, with two from one individual, so further material is required to assess
the variability of this feature.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

G. C. YOUNG 19

The ornament in C. pambulensis is much finer than in C. antarctica or C. stewarti.
Small triangular tubercles with posterodorsally directed points predominate in front of
the sensory canal, as in the other species, and behind the canal these tubercles become
more elongate and posteriorly directed (Fig. 2B, D), and may coalesce into subparallel
ridges (Figs 2C, 4B). Specimen CPC 17004 previously mentioned by Young (in Fergus-
son et al., 1979) is referred to C. pambulensis only on its similar ornament, since no sen-
sory grooves are preserved.

DISCUSSION

Morphology and Systematics

Denison (1979) included two genera in his family Diplacanthidae — Dizplacanthus
Agassiz and Gladiobranchus Bernacsek and Dineley. Long (1983a) preferred to place
Gladiobranchus as closely related to Uraniacanthus, as originally suggested by Bernacsek
and Dineley (1977). However Bernacsek and Dineley supported this relationship by
suggesting that Uraniacanthus may have possessed dermal plates and prepectoral spines,
even though Miles (1973) expressed no doubt from detailed study of the specimens that
these structures were absent in Uranzacanthus. Bernacsek and Dineley (1977) also inferred
that Gladiobranchus had dentigerous jawbones, but the holotype shows anterior circum-
orbital bones, indicating proximity of the snout, but no sign of ossified jaw elements.
These may have been lost, but the available evidence favours the interpretation that
Gladiobranchus did not possess them. Supporting this, and the climatiid affinities of
Gladiobranchus, are the presence of dermal shoulder plates, prepectoral spines, and the
large plates and tesserae of the head (Denison, 1979). The scapulocoracoid of Gladio-
branchus was restored by Bernacsek and Dineley (1977) as alow and broad element with a
truncated dorsal end, but the specimen (Bernacsek and Dineley, 1977: pl. 7) suggests a
higher narrow dorsal termination, with an anterior ridge and expanded posterior
ventral part, just as in Diplacanthus. The two enlarged circumorbitals, one at the
posterodorsal corner of the orbit, are special features shared with Dzplacanthus, as
Denison (1979) noted. Thus I follow Denison (1979) in considering Gladiobranchus better
placed as a diplacanthoid acanthodian, rather than an ischnacanthid (see also Long,
1986b: 335).

Long (1983a) modified Miles’s (1966) diagnosis of the Diplacanthoidei to include
Culmacanthus. The most significant resemblances (Fig. 5) are in the large dermal cheek
plate crossed by the infraorbital sensory groove, the relatively deep body, and the deeply
inserted spines. Culmacanthus differs from Diplacanthus primarily in the larger size of the
cheek plate, the absence of intermediate fin-spines, the probable absence of a mandi-
bular bone, the high narrow scapulocoracoid, and the structure of the dermal shoulder
girdle, with an unpaired lorical plate, and no spines attached to the paired pinnals. Long
(1983a) suggested that the cheek plate of Culmacanthus may have been homologous to the
postorbital or preopercular bones of Diplacanthus, but since there is no sensory groove
passing posterolaterally across the plate it seems more likely that it is only a much en-
larged postorbital, which extended more dorsally than the corresponding bone of
Diplacanthus, to include the junction of the posterodorsal branch of the infraorbital canal
at its ossification centre. For functional reasons it is likely that the cheek plate only
covered the region of the palatoquadrate, with a flexible hyoid gill cover situated
between it and the scapulocoracoid. The well-ossified branchiostegal rays in this
position in Gladiobranchus may be interpreted as a symplesiomorphy, by outgroup com-
parison with climatiids. It is unclear from Long’s (1986b) analysis of the dermal shoulder
girdle of climatiids whether the condition in Culmacanthus is primitive or specialized, but
if intermediate fin-spines and pectoral spines firmly attached to the pinnal plates are

PROC. LINN. SOC. N.S.W., 111 (1), 1989

20 DEVONIAN FOSSIL FISHES

Fig. 5. Culmacanthus stewartt Long. Restoration of complete fish, slightly modified after Long (1983a: fig. 9).

synapomorphies of ‘higher climatiids’ as he suggests (1986b: 337), then Dzplacanthus
would be more closely related to some Early Devonian climatiids than to Culmacanthus.
All students of acanthodian evolution have acknowledged the small number of reliable
characters available to define the major groups, and in the case of Culmacanthus the main
evidence still to be forthcoming concerns the histology of the scales and fin spines. This
is not known in the Victorian material because the bone is badly weathered. Better-
preserved material permitting histological study can be expected to clarify the relation-
ships of this form to the Euramerican diplacanthids.

Buostratigraphy

A biostratigraphic analysis of the Aztec fish fauna based on the distribution of
antiarchs was presented by Young (1988), in an attempt to resolve previous differences of
opinion about its precise age. Woodward (1921) first proposed a Late Devonian age on
the evidence of the antiarch Bothriolepis, and associated shark and large osteolepid scales,
and Gross (1950) reached the same conclusion. However White (1968) noted the
apparent mixing in the Aztec fauna of groups that elsewhere range in age from Middle
or even Early Devonian, to Late Devonian or Early Carboniferous, and on this basis he
favoured a late Middle Devonian rather than early Late Devonian age. Ritchie (1975)
added the placoderms Phyllolepis and Groenlandaspis to the faunal list, and proposed a
younger Late Devonian age. Young (1982) suggested correlations with southeastern
Australia based on the sharks in the Aztec fauna, which were consistent with Helby and
McElroy’s (1969) assessment of a Frasnian age for a palynoflora dominated by the spore
Geminospora lemurata. This species was assessed by Playford (1983) to have a known bio-
stratigraphic range from early or middle Givetian to late Frasnian or early Famennian.
However in Europe McGregor (1979) has estimated a much greater (Emsian-
Famennian) range for the possibly synonymous Gemuinospora svalbardiae, so the spore
evidence at present only dates the Aztec Siltstone within broad limits.

Other evidence supporting comparisons between the Aztec succession and the
Devonian sequences of southeastern Australia include the acanthodian Gyracanthides
described from Victoria by Woodward (1906), and from the upper Aztec succession by
White (1968), the antiarch Pambulaspis from southeastern New South Wales (Young,

PROC. LINN. SOC. N.S.W., 111 (1), 1989

G. C. YOUNG 21

1983) and described by Young (1988) from the top of the Aztec, and the culmacanthid
acanthodians just described. Culmacanthus antarctica 1s associated with turiniid thelodont
scales and bothriolepid antiarchs, and may therefore be the oldest known species, since
no thelodonts have been recorded from the successions in eastern Victoria and the south
coast of New South Wales. Young (1988) noted that the association of bothriolepid anti-
archs with thelodont scales is currently known from only one other locality, in the
Hatchery Creek fauna from near Wee Jasper, New South Wales (Young and Gorter,
1981). A post-Emsian age for this fauna is indicated by conodonts in the underlying
Murrumbidgee Group (serotznus Zone or slightly younger), but a younger age limit is un-
certain. In contrast the two Australian species of Culmacanthus are associated with
phyllolepid placoderms, which occur widely in southeastern Australia, in all other post-
Emsian faunas currently known which contain placoderms. From eastern Victoria and
the southern coast of New South Wales only the Bunga Beds fauna (Young, 1982) lacks
phyllolepids, but this is an impoverished fauna in which placoderms have not been
found. The abundant placoderms in the Aztec Siltstone includes phyllolepids only in the
highest horizons, and Young (1988) suggested that the base of the phyllolepid zone
preserved in the Aztec Siltstone, and the first appearance of the antiarch Pambulaspis,
may be older than the occurrence of related forms in eastern Victoria and southeastern
New South Wales.

In eastern Victoria, Long (1983b: fig. 3) used the appearance of Bothriolepis and
phyllolepids to define the base of the Upper Devonian, with Groenlandaspis occurring in
the latter half of the Frasnian. The fish fauna in the Boyd Volcanic Complex (Fergusson
et al., 1979; Young, 1982, 1983), which as well as Culmacanthus includes at least four taxa
(Bothriolepis, Pambulaspis, Antarctilamna, phyllolepids) in common with the Aztec fauna,
apparently lacks Groenlandaspis. The fact that both Bothriolepis and Groenlandaspis in the
Aztec Siltstone occur in the lowest fossiliferous beds, and far below the phyllolepid zone,
indicates again that these are earlier occurrences than the range of these forms in
southeastern Australia. In western New South Wales and central Australia groen-
landaspid arthrodires are associated with thelodont scales in the Wuttagoonaspis fauna
(Ritchie, 1975; Turner et a/., 1981) but the only antiarchs from this fauna are asterolepids
(Young, 1984a), so this appears to be older than the Aztec fauna. An Early-Middle
Devonian age has been assigned to the Wuttagoonaspis fauna, but the occurrence of the
Antarctic shark Mcmurdodus (Turner and Young, 1987), otherwise only known from the
Aztec Siltstone, may indicate less difference in age to the lower Aztec fauna than
previously thought.

In southeastern New South Wales the Boyd Volcanic Complex provides evidence
for a minimum age for the Aztec fauna, because the overlying Merrimbula Group con-
tains a marine invertebrate fauna of probable late Frasnian age (e.g. Roberts et al.,
1972). A single early Late Devonian marine transgression in southeastern Australia
may be assumed, for which the oldest reliable age is late Frasnian based on upper gigas
Zone conodonts (Pickett, 1972). This implies a late Givetian — early Frasnian age for the
underlying fishes and equivalent faunas in eastern Victoria, and an approximate
younger limit to the age of the Aztec Siltstone based on the phyllolepid placoderms in its
upper horizons. However, although the Victorian and southern New South Wales fish
successions are broadly equivalent in age, detailed taxonomic work on placoderms
common to the two areas (e.g. bothriolepids, phyllolepids) has not yet been done. The
results of the present study show specific differences between the Culmacanthus remains
from the two sequences, which may provisionally be regarded as indicating difference in
age, but there is as yet no reliable evidence on which occurrence 1s the older. It should
also be noted that according to Long (1983a) the culmacanthid from Freestone Creek is
not conspecific with C. stewartz, although other species (e.g. Bothriolepis cullodenensis Long,

PROC. LINN. SOG. N.S.W., 111 (1), 1989

22 DEVONIAN FOSSIL FISHES

1983b) are shared between the two assemblages. Further detailed taxonomic work on
the placoderms in the various faunas from southeastern New South Wales and Victoria
should clarify the precise correlation between the two sequences.

Biogeography

The biogeography of the Aztec fish fauna has been discussed previously by Young
(1981, 1982, 1984b, 1988) and Grande and Eastman (1986), whilst Long (1986a) has
recently commented on acanthodian biogeography in the Devonian. The presence of
species of Culmacanthus in southeastern Australia and southern Victoria Land, Antarc-
tica, but nowhere else, corresponds with the pattern indicated previously by the elasmo-
branch Antarctilamna (Young, 1982), the antiarch Pambulaspis (Young, 1988), and possibly
the placoderm Antarctaspis (if a relationship to phyllolepids is confirmed; see Young,
1987: fig. 5). These distributions are all consistent with grouping the two regions
together in an ‘East Gondwana Province’ as proposed by Young (1981). It should also be
noted that the only two fish assemblages recorded from any continent in which bothrio-
lepid antiarchs and turiniid thelodonts are associated are in southeastern Australia and
southern Victoria Land, and species group interrelationships for the cosmopolitan anti-
arch Bothriolepis again indicate an Antarctica-Australia connection (Young, 1988). All
these fishes inhabited freshwater environments, but this does not mean that they were
unable to cross marine barriers, and in this respect no specific data are provided on the
Devonian palaeogeography of the Antarctic-Australian region. It is relevant to note
however that the distinctive dermal cheek plates of Culmacanthus have not been found in
the well-studied fish faunas from the classic Old Red Sandstone localities of the North-
ern Hemisphere, but that they do occur in the Southern Hemisphere in two areas
separated today by the vast expanse of the Southern Ocean, and a distance in excess of
4500km, which exceeds by a considerable margin the 3000km-wide ocean postulated by
some authors to have separated Gondwana from Euramerica during the Late Devonian.
Thus the evidence of culmacanthid distribution is entirely consistent with reconstruc-
tions of a Palaeozoic Gondwana which minimize or remove oceanic barriers by juxta-
posing southeastern Australia and Victoria Land (Fig. 1).

ACKNOWLEDGEMENTS

The opportunity to participate in the Victoria University of Wellington Antarctic
Expedition of 1970-71, and the field and logistic support given to fossil collection in
southern Victoria Land, is gratefully acknowledged. R. W. Brown assisted in field
collection in the Eden-Pambula region, and took the photographs. W. Peters prepared
the specimens, and Dr John Long provided latex casts of Culmacanthus stewart: for com-
parison, and commented on the new specimens and the manuscript. Dr R. Ryburn
assisted with the bibliography. The paper is published with the permission of the
Director, Bureau of Mineral Resources, Canberra.

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McGReEGor, D. C., 1979. — Spores in Devonian stratigraphical correlation. Spec. Pap. Palaeontol. 23: 163-184.

McKe Lvey, B. C., Wess, P. N., GorTON, M. P., and KOHN, B. P., 1972. — Stratigraphy of the Beacon
Supergroup between the Olympus and Boomerang Ranges, Victoria Land. In Antarctic Geology and
Geophysics, R. J. ADIE (ed.): 345-352. Oslo: Universitets forlaget.

MILEs, R. S., 1966. — The acanthodian fishes of the Devonian Plattenkalk of the Paffrath trough in the
Rhineland. Ark. Zool. 18: 147-194.

—, 1973. — Articulated acanthodian fishes from the Old Red sandstone of England, with a review of the
structure and evolution of the acanthodian shoulder-girdle. Brit. Mus. (Nat. Host.) Bull. (Geol.) 24:
115-213.

PICKETT, J., 1972. — Late Devonian (Frasnian) conodonts from Ettrema, New South Wales. /. Proc. Roy. Soc.
N.S.W. 105: 31-37.

PLAYFORD, G., 1983. — The Devonian miospore genus Geminospora Balme, 1962: a reappraisal based on topo-
typic G. lemurata (type species). Mem. Ass. Australas. Paleont. 1: 311-325.

RitcHik, A., 1971a. — Ancient animals of Antarctica — Part 2. Hemisphere 15 (12): 12-17.

— , 1971b. — Fossil fish discoveries in Antarctica. Aust. Nat. Hist. 17: 65-71.

—., 1974. — From Greenland’s icy mountains. Aust. Nat. Hist. 18, 28-35.

——.,, 1975. — Groenlandaspis in Antarctica, Australia and Europe. Nature 254: 569-73.

ROBERTS, J:, JONES, P) J., JELL, J. S|, JENKINS, I. B: H., MARSDEN, M. A. H., MeKELLAR, R. G.,
McKELveY, B. C., and SEDDON, G., 1972. — Correlation of the Upper Devonian rocks of Australia.
J. geol. Soc. Aust. 18: 467-490.

TURNER, S., JONES, P. J., and DRaPER, J. J., 1981. — Early Devonian thelodonts (Agnatha) from the Toko
Syncline, western Oueensland, and a review of other Australian discoveries. BMR J. Aust. Geol.
Geophys. 6: 51-69.

, and YOUNG, G. C., 1987. — Shark teeth from the Early-Middle Devonian Cravens Peak Beds,
Georgina Basin, Queensland. Alcheringa 11: 233-244.

Watson, D. M. S., 1937. — The acanthodian fishes. Phil. Trans. Roy. Soc. Lond. 228: 49-146.

White, E. I., 1968. — Devonian fishes of the Mawson-Mulock area, Victoria Land, Antarctica. Scient. Rep.

transantarct. Exped. 16: 1-26.

PROG. LINN. SOG. N.S.W., 111 (1), 1989

24 DEVONIAN FOSSIL FISHES

WOODWARD, A. S., 1906. — On a Carboniferous fish fauna from the Mansfield district, Victoria. Mem.
National Mus., Melbourne 1: 1-32.

——, 1921. — Fish-remains from the Upper Old Red Sandstone of Granite Harbour, Antarctica. British
Antarctic (“lérra Nova’) Expedition, 1910. Nat. Hist. Rep. (Geol. ) 1: 51-62.

YOUNG, G. C., 1981. — Biogeography of Devonian vertebrates. Alcheringa 5: 225-243.

——, 1982. — Devonian sharks from south-eastern Australia and Antarctica. Palaeontology 25: 817-843.

— , 1983. — A new antiarchan fish (Placoderm1) from the Late Devonian of southeastern Australia. BMR
J. Aust. Geol. Geophys. 8: 71-81.

——, 1984a. — An asterolepidoid antiarch (placoderm fish) from the Early Devonian of the Georgina Basin,
central Australia. Alcheringa 8: 65-80.

——, 1984b. — Comments on the phylogeny and biogeography of antiarchs (Devonian placoderm fishes),
and the usc of fossils in biogeography. Proc. Linn. Soc. N.S.W. 107: 443-473.

—, 1987. — Devonian palaeontological data and the Armorica problem. Palaeogeog. Palaeoecol. Palaeoclimat.
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——, 1988. — Antiarchs (placoderm fishes) from the Devonian Aztec Siltstone, southern Victoria Land,

Antarctica. Palaeontographica A202: 1-125.
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YOUNG, V. T., 1986. — Early Devonian fish material from the Horlick Formation, Ohio Range Antarctica.
Alcheringa 10: 35-44

PROC. LINN. SOC. N.S.W., 111 (1), 1989

A new Australian Species of Argyrodes
Simon (Araneoidea: ‘Theridiidae)
which preys on its Host

M. R. GRAY and G. J. ANDERSON

Gray, M. R., & ANDERSON, G. J. A new Australian species of Argyrodes Simon
(Araneoidea: Theridiidae) which preys on its host. Proc. Linn. Soc. N.S.W. 111 (1),
1989: 25-30.

A new species of the web-parasite spider genus A7gyrodes is described. This spider
has a close association with its only known and larger host, the tent web spider
Achaearanea mundula. It is a predator of both the host and its eggs and young and makes
use of the host’s leaf retreat as a shelter for its egg sac.

M. R. Gray, The Australian Museum, PO. Box A285, Sydney South, Australia 2000, and G. J.
Anderson, Dept of Medicine, University of Queensland, Royal Brisbane Hospital, Herston, Aus-
tralia 4029; manuscript received 5 January 1988, accepted for publication 19 October 1988.

INTRODUCTION

Large, complex or communal spider webs may harbour a variety of invertebrates as
temporary or semi-permanent residents. The small spiders of the worldwide genus
Argyrodes Simon (Exline and Levi, 1962) are common residents in the webs of araneoid
spiders. Most live either as commensals which take small prey ignored by the host spider
or kleptoparasites which steal prey captured by the host (Robinson and Robinson, 1973;
Vollrath, 1979; Whitehouse, 1986). The host spider is often much larger (e.g. Nephila or
Argiope spp.) than the resident Argyrodes so that competition for food resources would not
seem likely. Several studies have shown, however, that prey stealing can have such an
impact that some hosts abandon their webs and build elsewhere (Robinson and Robin-
son, 1976; Vollrath, 1981; Rypstra, 1981). A few temperate zone species of Argyrodes have
become predators of both their hosts and spiderlings (Kaston, 1965). Trail (1980) has
observed host predation by A. fictzlium (Hentz) and A. baboquivari Exline and Levi upon
linyphiid host spiders including Frontinella sp., and the uloborid Philliponella owent
Chamberlin. A. trigonum (Hentz) is a predator of the araneids Metepeira labyrintha Hentz
and Mecynogea lemniscata Walckenaer and the linyphiid Nerene radiata Walckenaer (Wise,
1982; Larcher and Wise, 1985). Whitehouse (1986) noted predation by A. antipodiana
O.P. — Cambridge upon its araneid host Avaneus pustulosa Walckenaer while the host was
moulting and virtually defenceless. Gray (1983) briefly reported upon a distinctive
Argyrodes species from eastern Australia which showed host predation behaviour. A
description of this spider and some behavioural observations are given below.

Argyrodes incursus new species
Figs 4-12
Types: Holotype male (KS 18359), Blackbutt Reserve, Newcastle, New South Wales,
March 1986, G. Anderson. Paratypes: Allotype female (KS 18360), 3 females (KS
18361), data as above; 2 females (KS 12254-55), Upper Causeway and Couranga Track,
Royal National Park, near Sydney, New South Wales, 4.3.1983, M. Gray and C. Horse-
man. Australian Museum coll.

Material examined: 2 females, Station 32, southwest of North Hummock, Lord Howe
Island, New South Wales, 6.2.1971, M. Gray.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

26 HOST-PREDATING ARGYRODES

Figs 1-3. 1, Argyrodes incursus, ‘engorged’ female. 2-3, Achaearanea mundula: 2, female; 3, web. Scale lines: Figs 1-
2, 1.0mm.

Diagnosis. Male palp with marginal coiled embolus, abdomen with dorsal scute.
Female with orange-red spot on posterodorsal abdomen, abdomen rounded.

Male. Total length 2.18. Carapace length 0.90, width 0.63. Abdomen length 1.27, width
0.89. Leg 1 segment lengths: femur 1.07, patella 0.33, tibia 0.63, metatarsus 0.65, tarsus
0.45. Carapace, sternum and mouthparts dark brown. Abdomen dark brown dorsally
and basally; remainder black apart from 4 small orange spots placed around the base of
the spinnerets (Figs 4, 5). Femora and tibiae of legs 1 and 2 dark brown, femora 2 lighter

PROC. LINN. SOG. N.S.W., 111 (1), 1989

M. R. GRAY ANDG. J. ANDERSON 27

ventrally; remaining segments reddish brown. Legs 3 and 4 reddish brown, femora
lightest. Cheliceral groove with 3 promarginal, 1 retromarginal teeth. A.M.E. largest,
other eyes subequal. A.M.E. separated from each other by almost 2 diameters and from
A.L.E. by 0.5 diameters; P.M.E. separated by 1.75 diameters. Clypeus with wide, trans-
verse groove with numerous anterodorsally directed setae in and below groove. Trans-
verse stridulatory ridges run across the posterolateral carapace lobes. Abdomen ovoid,
widest behind middle and covered with a dorsal scute which extends anteriorly around
the pedicel. A separate scute surrounds the spinnerets (Fig. 5). Dorsal pedicel margin
strongly sclerotised with evenly spaced enlarged hair bases as stridulatory picks. Male
palp (Fig. 11) with tightly coiled embolus (3 turns) surrounding a raised central tegular
area. Embolus supported marginally by a membranous conductor. Radix apical, tri-
angular and lying adjacent to the elongate median apophysis.
Female. Total length 3.36-4.33. Carapace length 1.29-1.35, width 0.88-0.92. Abdomen
length 1.97-3.00, width 1.80-2.55. Leg 1 segment lengths: femur 1.20-1.25, patella 0.35-
‘0.38, tibia 0.80-0.96, metatarsus 0.80-0.88, tarsus 0.50-0.62. Carapace, sternum and
mouthparts dark brown. Legs 1-3 dark brown except metatarsi and tarsi which are
reddish brown; leg 4 reddish brown. Abdomen ovoid to spherical, black with two in-
distinct rows of unpigmented spots anterodorsally. Posterodorsal abdomen with a large
orange-red spot (occasionally divided into two unequal spots), bounded anteriorly and
posteriorly by 1 or 2 narrow transverse orange bars (Figs 7, 8, 9). Four small irregular
orange spots are grouped around the spinnerets. Epigynum (Fig. 6) a low mound sur-
mounted by two ovoid fossae separated by about half a diameter. Internal genitalia (Fig.
10) with more or less spherical receptacula separated by about half their width. Sperm
ducts strongly convoluted.

+ MORPHOLOGY AND BEHAVIOUR

Both the shape and coloration of the abdomen of Argyrodes incursus are unusual.
Elongation, shoulder or posterior tubercles and silvery ornamentation are common
features of the Argyrodes abdomen (Exline and Levi, 1962). In unfed female specimens of
A. incursus the abdomen is ovoid but expands considerably after feeding into a smooth
sphere (Fig. 1). A prominent orange-red spot (absent in males) on the posterodorsal
abdomen contrasts markedly with the shiny black coloration of the remainder. The
function of this distinctive female colour pattern is not understood but may have some
aposematic benefit, such as providing protection from other web-invading predators
(e.g. mimetid spiders or wasps). A dark brown dorsal scute covers the male abdomen.
The male palp is unusual in that the tegulum forms a central ‘drum’ around which the
coiled embolus is wound.

A. incursus was always found in association with the ‘tent web’ spider, Achaearanea
mundula (LL. Koch). This spider (Fig. 2) is common in open forest habitats in eastern
New South Wales and Queensland. Populations with which A. zncursus was associated
were located at Newcastle, Sydney, and Lord Howe Island, New South Wales. Ach.
mundula builds a complex web consisting of a knockdown maze of threads above a finely
woven horizontal sheet (Fig. 3). The curled leaf or litter retreat is placed at the centre of
the knockdown maze; this provides a refuge for the spider and its egg sacs (up to 6). The
webs are usually built among low understorey foliage. They also harbour commensal/
kleptoparasitic species of Argyrodes plus uloborid and mimetid spiders; the latter are
temporary residents, well known as predators of araneoid spiders.

Specimens of A. incursus were found singly inside the curled leaf retreats of Ach.
mundula individuals during February and March. In the Newcastle population, however,
the catching parts of 3 host webs harboured 8, 2 and 2 A. incursus individuals respec-

PROG. LINN. SOG. N.S.W., 111 (1), 1989

28 HOST-PREDATING ARGYRODES

>
3; EX

Figs 4-12. Argyrodes incursus. 4-5, male body: 4, dorsal; 5, lateral. 6, epigynum. 7-9, female abdomen: 7,
dorsal; 8, lateral; 9, ventral. 10, female internal genitalia, dorsal. 11, male palp, ventral. 12, upper part of
curled leaf retreat of Ach. mundula split open to show A. incursus egg sac near apex. Scale lines: Figs 4-11,
0.5mm; Fig. 12, 3.5mm.

PROC. LINN. SOC. N.SW., 111 (1), 1989

M. R. GRAY ANDG. J. ANDERSON 29

tively, with the female hosts still occupying their retreats. Six observations were made of
A. incursus females actually feeding upon the carcass of their host spider and/or its eggs or
spiderlings, always within the leaf retreat. The dead host spiders were partially silk
wrapped and their loosely woven egg sacs, if attacked, were torn open. In the Newcastle
population A. zncursus individuals also were found living inside the leaf retreats of empty
Ach. mundula webs; in one case a male and female were found together. Some of these
females had built single egg sacs at the leaf retreat apices (Fig. 12). Seven egg sacs were
observed, all with the characteristic urn shape found in many species of Argyrodes (Exline
and Levi, 1962). When present, the female sat above its sac on the stalk attachment.

At present Ach. mundula is the only known prey of A. zncursus. The capacity for
almost tick-like engorgement of the abdomen of A. incursus after feeding is particularly
striking. Females may be able to survive and reproduce on the food reserves obtained
from one successful predatory episode. The limited observations suggest that mating
may occur within the Ach. mundula retreat. In addition, A. incursus females use the empty
host leaf retreats for the protected placement of their egg sacs. All of this suggests a close,
perhaps exclusive, association of A. zncursus with its host/prey species, Ach. mundula.

In a study of A. trigonum (Hentz), Larcher and Wise (1985) observed a wide range of
behaviour including commensalism, kleptoparasitism, web theft and host predation.
They concluded that the nature of the interaction between A. trigonum and its hosts
varied as a function of their relative individual sizes. This agrees with an earlier finding
of Trail (1980) that Argyrodes species which attack their hosts are equal or larger in adult
size compared with their host species. Presumably, this makes the attacker better able to
Overpower its prey. A. incursus, however is considerably smaller than its prey, Ach.
mundula. Carapace length ranges for females of A. zncursus and Ach. mundula are respec-
tively 1.2971.35mm (M = 1.32) and 2.17-2.58mm (M = 2.32). The mean difference of
1.00mm is highly significant (P< 0.001). The suggestion of Trail (1980) that an Argyrodes
species smaller than its host would behave as a commensal or kleptoparasite rather than
as a host predator is not supported in this case.

As yet, little information about the attack behaviour of A. incursus is available. It
seems likely that the host is attacked in its retreat as all carcasses were found in retreats.
Limited observations by Larcher and Wise (1985) suggest that A. trigonum simply
approaches the host and bites it on a leg, resistance being negligible (though some hosts
flee to avoid an attack). The venom is apparently quick to immobilise the prey which 1s
wrapped after being bitten. An intriguing but anecdotal observation (Walker, 1983)
notes that a spider purported to be an A7gyrodes species entered the web of a red-back
spider (Latrodectus hasselti Thorell) and squirted a milky substance (source unknown)
onto the silk. The red-back readily ate the substance and became immobilized soon
after. Such an indirect prey capture technique would provide an effective predation
strategy in a small predator/larger prey system. Alternative hypotheses include stealth,
the prey remaining unaware of the slowly approaching predator; or male behavioural
mimicry, the predator imitating some aspect of the courtship repertoire of the prey’s
male.

The level and impact of A. incursus predation upon Ach. mundula populations are not
clear. Data for a small sample of 17 Ach. mundula webs from Royal National Park near
Sydney showed that only 2 webs harboured A. incursus individuals. By contrast 13 webs
contained other presumptively kleptoparasitic/commensal A7gyrodes residents. However,
Newcastle Ach. mundula webs showed A. incursus occupancy rates of approximately 50%.
This suggests that A. zncursus predation may exert a considerable effect at least in small
host populations.

PROG. LINN. SOG. N.S.W., 111 (1), 1989

30 HOST-PREDATING ARGYRODES

ACKNOWLEDGEMENTS

We thank Dr H. W. Levi, Museum of Comparative Zoology, Harvard University,
for confirming the generic diagnosis.

References

EXLINE, H., and Levi, H. W., 1962. — American spiders of the genus Argyrodes. Bull. Mus. Comp. Zool. 127:
75-204.
Gray, M. R., 1983. — Spider predation on Achaearanea mundula L. Koch. Aust. Arachnol. 13: 6-8.
KASTON, B. J., 1965. — Some little known aspects of spider behaviour. Amer. Midl. Nat. 73: 336-356.
LARCHER, S. F., and WIsE, D. H., 1985. — Experimental studies of the interaction between a web-invading
spider and two host species. J. Arachnol. 13: 43-59.
ROBINSON, M. H., and ROBINSON, B., 1973. — Ecology and behaviour of the giant wood spider Nephila
maculata (Fabr.) in New Guinca. Smithson. Contr. Zool. 149: 1-76.
, and , 1976. — The ecology and behaviour of Nephila maculata: a supplement. Smithson. Contr. Zool.
218: 1-22.
RypsTRA, A. L., 1981. — The effect of kleptoparasites on prey consumption and web relocation in a Peruvian
population of the spider Nephila clavipes (L.) (Arancae: Araneidae). Ozkos 37: 179-182.
TRAIL, D. S., 1980. — Predation by Argyrodes (Theridiidae) on solitary and communal spiders. Psyche 87:
349-355.

VOLLRATH, F., 1979. — Behaviour of the kleptoparasitic spider Argyrodes elevatus (Araneac, Theridiidac).
Anim. Behav. 27: 515-521.

——, 1981. — Energetic considerations of a spider parasite-spider host system. Rev. Azachnol. 3: 37-44.

WALKER, P., 1983. — Note in: Aust. Arachnol. 14: 9.

WHITEHOUSE, M. E. A., 1986. — The foraging behaviours of Argyrodes antipodiana (Theridiidac), a klepto-
parasitic spider from New Zealand. New Zealand J. Zool. 13: 151-168.

Wise, D. H., 1982. — Predation by a commensal spider, Argyrodes trigonum, upon its host: an experimental
study. J. Arachnol. 10: 111-116.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

Two new Species of Amphientomidae
(Insecta: Psocoptera), the first Record of the
Family for Australia

C. N. SMITHERS

SMITHERS, C. N. Two new species of Amphientomidae (Insecta: Psocoptera), the first
record of the family for Australia. Proc. Linn. Soc. N.S.W. 111 (1), 1989: 31-35.

Two new species of Amphientomidae, representing the first species of the family to
be recorded from Australia, are described.

C. N. Smithers, Research Associate, Australian Museum, College St., Sydney, Australia 2000;
manuscript received 22 June 1988, accepted for publication 24 August 1988.

INTRODUCTION

The psocopteran family Amphientomidae has not been recorded from Australia
although Mockford (in lit.) has mentioned that the family occurs here. On this authority
the family was noted as being Australian in the manuscript of the forthcoming revised
edition of the ‘Insects of Australia. While sorting material in the collections of the
Australian Museum two specimens have been found, each belonging to an undescribed
species. They are described here.

The Amphientomidae now include nearly 80 species, grouped into 18 genera. In
the main they are infrequently encountered insects of warm climates. Three genera are
known only from amber and there are a few species in the genus Amphientomum Pictet
which are also known from amber. Current generic definitions are based mainly on
venational features but these are gradually being supplemented by other characters as
additional material becomes available.

The Amphientomidae are unusual in that they are one of only two families in the
order in which the body is clothed with scales, the other family being the unrelated
Lepidopsocidae. The members of both bear a superficial resemblance to micro-
lepidopterans and can easily be mistaken for them in the field.

Definitions and illustrations of the main distinguishing features of the family and
genera have been given in Smithers(1972); they are not repeated here. The two new
species fall clearly into the genera Hemiseopsis Enderlein and Seopsts Enderlein
respectively.

SYSTEMATIC DESCRIPTIONS
Hemuseopsis alettae sp. nov.

FEMALE
Coloration (in alcohol). Head and body pale testaceous, the head a little darker than
body and legs. Eyes reddish brown.

Morphology. Length of body: 3.1mm. Median epicranial suture fine but very dis-
tinct, anterior arms absent. Pilosity of head very fine. No hairs between ommatidia.
Length of flagellar segments: f,: 0.14mm; f: 0.19mm. First flagellar segment broader in
basal half than in distal half, attachment to pedicel very narrow. Second flagellar seg-
ment longer than first. Antennal striations very fine and dense. Scape and pedicel
broad. Eyes large, not protruding from head capsule except behind. Dorsal margin con-
tinuous with vertex. Seen from above eyes curve towards each other anteriorly continu-
ing curvature of head. IO/D: 1.6; PO: 0.5. Lateral ocelli small, situated about a third of

PROC. LINN. SOC. N.S.W., 111 (1), 1989

ae NEW SPECIES OF AMPHIENTOMIDAE (INSECTA PSOCOPTERA)

distance from eye to median epicranial suture, opposite aboyt middle of eye. Lacinia
(Fig. 2) curved. Maxillary palp densely beset with microtrichia. Second segment with
small sensory cone and broader than third and fourth segments. Measurements of hind
leg: F: 0.6mm; T: 1.0mm: t;: 0.68mm: ty: 0.11mm; ts: 0.13mm; rt: 6.2:1:1.2; ct:27,0,0.
Ctenidiobothria very strong with well developed basal combs. Femur of anterior legs
without cones along internal margin. Tibia of second pair of legs with external spine
about half way along length in addition to usual apical spines. Femur of third leg with
strong, curved, dorsal apical spine. Tibia of third leg with seven ventral spines in distal
half in addition to usual apical spines. Claws with one preapical tooth and row of setulae
basad of tooth. Fore wing length: 2.5mm; width: 0.78mm. Fore wing (Fig. 1) almost
parallel-sided, rounded apically. Basal section of Sc ends in somewhat thickened costa.
Distal section of Sc absent. R, parallel with costal margin basad of the elongate stigma-
pophysis beyond which it bends to meet costal margin. Rs divides between origins of My
and M3. Cu, divides opposite stigmapophysis. Hind wing with R, present. M simple.
Epiproct lightly sclerotized, simply rounded behind with preapical row of setae basad of
which setae are sparsely and irregularly arranged. Paraprocts simple, without clearly
defined trichobothrial field but with a few large setae without patterned area around
base. Subgenital plate simple. Gonapophyses (Fig. 3) with ventral valve greatly attenu-
ated ending in sharp point, with ventral membranous flange. Dorsal valve broad in
basal half, narrowing abruptly to form long tapering extension which matches and lies
adjacent to that of ventral valve. External valve tapering to bluntly rounded end, with
strongly developed dorsal lobe.

MATERIAL EXAMINED. Holotype female, Falbrook, N. Singleton, New South Wales,
26.1.1979. A. S. Smithers. Holotype in Australian Museum. This species is named for
my wife who collected the type specimen.

DISCUSSION. Four species of Hemiseopsis have now been described, three from Africa and
the present species from Australia. The genus may also occur in Jamaica (Turner, 1975).
H1. alettae is smaller than H. fuellerborni (Enderlein) from Tanzania. Zaire, Angola and
Zambia. In that species Rs divides opposite the origin of M3, Rs lies much further from
R, and Cu, is much shorter than in H. alettae. The basal section of Rs is very much
shorter than Ro,3. The genitalia of H. fuellerborni have not been described. H. machadoi
Badonnel, from Angola, is only a little larger than H. alettae but can be distinguished by
features of the wing venation and the female genitalia. In H. machadoi R, does not reach
the wing margin and the crossvein from R, meets M basad of its separation from Rs.
Ms arises opposite the division of Rs and Cu), is much shorter than in H. alettae. The
ventral and dorsal valves of the gonapophyses are not strongly attenuated, reaching only
as far as the end of the external valve. In H. alettae they extend far beyond the end of the
external valve. In H. machadoi the lacinia is straight with a different arrangement of
apical teeth from that in H. alettae. In H. obscurus Broadhead and Richards, from Kenya,
several of the veins of the fore wing in the basal half are weakly differentiated. The
ventral and dorsal valves of the gonapophyses are not attenuated but are blunt; the
dorsal lobe of the external valve is longer than the valve itself, a condition not found in
any of the species of which the genitalia have been described. The head of H. obscurus has
a distinctive colour pattern not found in any of the other species.

Seopsis incisa sp. nov.

MALE
Coloration (in alcohol). Head pale brown with brown markings. A narrow brown

PROC. LINN. SOC. N.S.W., 111 (1), 1989

C. N. SMITHERS 33

“Imm

Figs 1-9. Hemiseopsis alettae sp. nov. 1, female fore wing. 2, female lacinia. 3, female gonapophyses. Seopsts incisa
sp. nov. 4, male femoral spines. 5, male lacinia. 6, male phallosome. 7, male fore wing. 8, male hind wing. 9,
male paraproct.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

34 NEW SPECIES OF AMPHIENTOMIDAE (INSECTA PSOCOPTERA)

mark on each epicranial plate along the ridge of the vertex; a diffuse, broad brown band
across the front of the head from eye to eye; a narrow band in position usually occupied
by anterior arms of epicranial suture, this broadened laterally to form a small patch just
anterior to the lateral ocellus on each side; posterior half of postclypeus with a reticulate
pattern in brown, part of which consists of narrow, longitudinal, parallel lines, the
pattern extending further anteriorly in the midline than laterally: postclypeus pale in
anterior half. Labrum pale. Lower half of genae brown. Antennae dark brown. Eyes
black. Ocelli circled with dark brown. Thorax pale brown, darker along some sutures.
Abdomen pale, darker dorsally on basal quarter and with suggestion of irregular seg-
mental marks, laterally dark. Femora pale. Fore and middle tibiae pale in basal half,
brown in distal half. Hind tibiae pale. First tarsal segment pale in basal half, darker in
distal half but with pale tip. Second and third segments brown. Fore wing membrane
hyaline, faintly tinged with brown, paler in distal third. Hind wing hyaline.

Morphology. Length of body: 2.3mm. Median epicranial suture fine but distinct.
Anterior arms indistinct but position marked by brown band. Vertex slightly curved.
Length of flagellar segments: f;:0.31m; f):0.34mm. Antennal flagellum very fine, with
very long setae from 7 to 15 times as long as flagellar width. Transverse striations of
antennae very numerous. Eyes large, but hardly protruding. Upper margin almost level
with vertex, extending a little behind head laterally. Fine, short setae between facets.
Three ocelli. Lateral ocelli close to antero-medial angle of eyes, median ocellus in
middle of frons so that the three ocelli are almost in a straight line. Lacinia (Fig. 5)
almost straight, deeply divided at apex. Measurements of hind leg: F: 0.5mm, T:
0.98mm; t,: 0.54mm; to: 0.l1mm; t3: 0.1mm; rt: 5.4:1:1; ct: 19,0,0. Front femur with long
row of short spines (Fig. 4). Claws with one preapical tooth. Fore wing length: 2.5mm;
width: 0.76mm. Fore wing (Fig. 7) narrows to somewhat pointed apex. Venation as
usual in genus but distal section of Sc and Cu), evanescent and less obvious than other
veins. Hind wing length: 2.0mm; width: 0.6mm. Hind wing (Fig. 8) with R,; not reach-
ing wing margin. IA curves distally towards hind margin. Epiproct simple, rounded
behind, setose. Paraproct (Fig. 9) with finely and densely rugose, slightly raised area
from which large setae arise. Bases of setae not surrounded by ‘rosette’ pattern often en-
countered in barklice. Hypandrium well sclerotized, simple, posteriorly and laterally
slightly upturned. Eighth sternite more heavily sclerotized than more anterior sternites
and forming an anterior extension to the hypandrium. Phallosome (Fig. 6).

MATERIAL EXAMINED. Holotype male, Malaise trap, AOS 125°50’E, mining
camp, Mitchell Plateau, Western Australia, 9-19.v.1983. I. D. Naumann and J. C.
Cardale. Holotype in Australian National Insect Collection, Canberra.

DISCUSSION. Seopsis is now a genus of ten species. There are three from Sri Lanka, two
from the Philippines, two from Angola and one from each of Nepal, Singapore and Aus-
tralia. S. brunnea New, from Singapore, differs from S. incisa in lacking the dark post-
clypeal pattern, in having a lacinia without a deeply divided apex and in lacking the row
of spines on the fore femur. The male phallosome is similar to that of S. incisa in general
form but differs in proportions. S. /uzonica Banks, from the Philippines, differs in having
a brown head with a pale transverse band at the level of the ocelli and in having two dark
marks on each side on the vertex. In S. metallops Enderlein, from Sri Lanka, the head is
black or dark brown with a pale spot in the middle of the front of the head and a similar
spot on each half of the vertex. The lateral ocelli are much further from the eyes than in
S. incisa. The membrane of the fore wing is dark. The basal segment of the hind tarsus is
more than seven times the length of the second segment, whereas it is only 5.4 times as
long in S. zncisa. S. nepalensis New, from Nepal, has the posterior part of the postclypeus

PROC. LINN. SOC. N.S.W., 111 (1), 1989

C.N. SMITHERS 35

pale, not dark as in S. zncisa. The fore wing 1s acuminate and there are accessory sclerites
internally at the posterior ends of the male parameres. The apex of the lacinia is not
deeply divided. S. pavonia Badonnel, from Angola, has the fore wing strongly acuminate
and the lacinia is not deeply divided. S. termitophila Badonnel, also from Angola, has
much shorter wings than S. znczsa (1.7mm as opposed to 2.5mm). The apex of the lacinia
is not deeply divided and the male phallosome has a pair of pitted plates, internally,
adjacent to the ends of the parameres. S. superba Hagen, from Sri Lanka, has a brown
head with a darker longitudinal band on the vertex. The ocelli are close togther. The
basal tarsal segment of the hind legs is very much longer than in S. incisa, being about
seven times as long as the second. S. trecolor Banks, from the Philippines, has a brown
head and the lateral ocelli are set well away from the eyes. It is larger than S. incisa (fore
wing length 3.5mm as opposed to 2.5mm). The illustration of the wing which accom-
panies the description shows no distal section to Sc, an unusual condition for this genus.
S. vasantasena Enderlein, from Sri Lanka, has a brownish yellow head with four narrow,
longitudinal streaks from the vertex down onto the front of the head; the postclypeus has
a double, broad brown longitudinal band. The fore wing length is only 2.0mm.

References

SMITHERS, GC. N., 1972. — The classification and phylogeny of the Psocoptera. Aust. Mus. Mem. 14: 1-349.
TURNER, B. D., 1975. — The Psocoptera of Jamaica. Trans Roy. ent. Soc. Lond. 126(4): 533-609.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

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Redistribution of Amino Acids and Amides

during Seedling Development in Acacia iteaphylla
F. Muell. (Fabaceae: Mimosoideae)

ANITA B. BRADKE and DAVID R. MURRAY

BRADKE, ANITA B., & MuRRAY, Davib R. Redistribution of amino acids and amides
during seedling development in Acacia iteaphylla F. Muell. (Fabaccae:
Mimosoideac). Proc. Linn. Soc. N.S.W. 111 (1), 1989: 37-42.

The distribution of free amino compounds in the cotyledons and the organs
derived from the embryonic axis has been studied in developing seedlings of Acacia
iteaphylla. The results indicate that asparagine and pipecolic acid are the major forms of
nitrogen translocated initially from the cotyledons to the hypocotyl and primary root.
Selective transport of several ‘non-protcin’ amino acids was indicated. S-carboxycthyl-
cysteine became prominent in the hypocotyl after the pipecolic acid content of the
hypocotyl had declined; it appeared also in the primary leaf, but was not strongly
represented in the roots. S-carboxyisopropylcysteine appeared late in the hypocotyl,
and was not detected in any other part of the axis. Albizziine was not transported out of
the cotyledons, but like arginine was metabolized zn situ.

Anita B. Bradke and David R. Murray, Biology Department, University of Wollongong, Wollon-
gong, Australia 2500 (private address D.R.M., 7 Acacia Avenue, Gwynneville, Australia 2500);
manuscript received 14 June 1988, accepted for publication 24 August 1988.

INTRODUCTION

The cotyledons of Acacia seedlings undergo a transition from storage organs to
photosynthetic organs (Ashcroft and Murray, 1979; Murray, 1981; Smith, 1981). During
this transition, reserve materials appear to be mobilized in two stages. The first encom-
passes the initial growth of the radicle and hypocotyl, resulting in the elevation of the
cotyledons. In Acacia iteaphylla F. Muell., the dry matter content of the cotyledons is
depleted by about 70% during this stage, but only a slight net decline in the protein con-
tent of the cotyledons is observed (Ashcroft and Murray, 1979). In the second phase,
more rapid net breakdown of protein in the cotyledons 1s associated with the synthesis of
chloroplasts, as the cotyledons and then the primary leaf become effective photo-
synthetic organs.

It has long been known that Acacia seeds contain a high proportion of total seed
nitrogen as free amino acids (Petrie, 1908, 1911). Many ‘non-protein’ amino acids have
now been identified (Seneviratne and Fowden, 1968), and their distribution among
species has indicated that there are four main sub-groups of Acacia throughout the world
(Evans e¢ al., 1977; Murray, 1986b). In view of the early slow rate of net decline in the
protein content of cotyledons, Ashcroft and Murray (1979) suggested that non-protein
amino acids might represent the earliest mobilized form of nitrogenous reserve.
Evidence has now been obtained that some, but not all, of the non-protein amino acids
are transmitted to the axis, together with newly-synthesized asparagine.

MATERIALS AND METHODS

The seeds of Acacia iteaphylla were from the same batch studied previously (Ashcroft
and Murray, 1979). To permit uniform imbibition, the seedcoats were cut with a razor
blade at the end furthest from the embryonic axis. Except for seeds imbibed for 24h,
which were placed in Petri dishes between moist Whatman No. 1 filter paper at 23°C
(Krishna and Murray, 1988), seeds were placed in trays of sandy soil in a glass house,

PROC. LINN. SOC. N.S.W., 111 (1), 1989

38 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS

with day temperatures around 30°C. At intervals up to 14 days, groups of 10 to 20
uniform seedlings were removed, dismembered and analysed.

Extracts from cotyledons, radicles and hypocotyls were prepared using a chilled
mortar and pestle, acid-washed sand, and medium consisting of 235mM K* phosphate
(pH 7.5), 0.5mM 2-mercaptoethanol and 0.03% (w/v) Triton X-100 (Murray and
Kennedy, 1980; Murray, 1983). The ratio of medium to tissue was 5:1 (mL per g fresh
weight). The homogenates were centrifuged for 4min at 9,000g in a Microfuge and the
supernatants removed. These were treated with ethanol (4:1, v/v) and insoluble material
removed by centrifugation (Murray and McGee, 1986). Smaller samples (whole axes;
balance of shoot) were extracted directly with ethanol (4:1, v/v). Aliquots of the ethanol-
soluble fractions were assayed for amino nitrogen content by the procedure of Schnar-
renberger et al. (1972), using L-serine as a reference standard (Murray, 1983).

Ethanol-soluble fractions were dried in a rotary evaporator, then redissolved in 0.5
to 1.0mL of 70% (v/v) ethanol, and applied to the origins of Whatman No. | papers pre-
pared for 2-dimensional descending chromatography (Murray, 1983). Extracts avail-
able only in small quantities were applied without prior concentration. The solvents
employed were 80% (w/v) phenol-water plus ammonia (200:1, v/v) in the first
dimension, then either n-butanol: acetic acid: water 12:3:5 (v/v) or n-butanol: propionic
acid: water 6:3:4 (v/v) in the second (Murray e¢ al., 1971; Murray, 1983). At least two
chromatograms were run for each extract. Amino compounds were detected by their
reaction with ninhydrin, and identified by comparison of their positions with those of
authentic compounds. Information on the chromatographic behaviour of non-protein
amino acids was kindly provided by Dr C. S. Evans and Prof. E. A. Bell. Authentic
albizziine was purchased from Aldrich; other amino acids and amides were from Sigma

Chemical Co.

RESULTS

The rate of seedling growth was faster than in the previous study because of higher
temperatures and longer day-length. The 4-day, 8-day and 14-day stages chosen for
analysis closely resembled the 8-day, 15-day and > 21-day stages described previously
(Ashcroft and Murray, 1979; Murray, 1981). At the 1-day stage, the amino nitrogen con-
tent of the cotyledons effectively represents that of the whole embryo (Fig. 1), as the axis
was too small for accurate analysis (less than Img fresh weight). The amount of free
amino nitrogen in the whole seedling progressively increased, doubling by 14 days (Fig.
1). Within the cotyledons, the amino nitrogen content did not alter substantially until
between 8 and 14 days, when it declined. Within the axis, the hypocotyl gained a much
higher proportion of the amino nitrogen exported by the cotyledons than did the radicle.

The amino acids and amides present in Acacia seedling tissues were identified as
shown in Fig. 2. Changes in their relative abundance in the cotyledons are shown in
Table 1. Many of the major nitrogenous solutes stored in the cotyledons initially are still
among the most prominent forms present after 14 days, when the total free amino nitro-
gen content of the cotyledons had declined by 40% (Fig. 1). The content of asparagine
increased, while the contents of albizziine, glycine, arginine, pipecolic acid, and finally
glutamate and alanine, declined (Table 1).

The distribution of amino compounds in the roots and hypocotyls of seedlings aged
8 and 14 days is shown in Table 2. Pipecolic acid and asparagine were the most abundant
forms of free amino nitrogen in both the roots and the hypocotyl] of the 8-day-old seed-
lings. At this stage, S-carboxyethylcysteine was prominent in the hypocotyl, but was not
detectable in the roots. At the 14-day stage, both S-carboxyethylcysteine and S-carboxy-
isopropylcysteine became prominent in the hypocotyl. The content of asparagine was

PROC. LINN. SOC. N.S.W., 111 (1), 1989

ANITA B. BRADKE AND DAVID R. MURRAY 39

Mmoles amino N
oOo + NY OO bh HO

See
A R

DAYS: 1 4 8 14

Fig. 1. Changes in the distribution of free amino nitrogen in the developing seedling of Acacia iteaphylla. C =
cotyledon pair; A = whole embryonic axis; R = radicle or root system; H = hypocotyl; S = shoot system
above the cotyledons.

NOISNIWIG puZg

PIPECOLIC
ACID

= ana

(ae

ORIGIN 1st DIMENSION

Fig. 2. The positions and identities of amino compounds separated from cotyledons of Acacia iteaphylla by 2-
dimensional descending paper chromatography. Solvent 1, phenol-water-ammonia; solvent 2, n-butanol-
acetic acid-water (sec Materials and Mcthods). Abbreviations are standard, plus S-CEC = S-

carboxyethylcysteine; S-CIC = S-carboxyisopropylcysteine.
; PROC. LINN. SOC. N.S.W,, 111 (1), 1989

40 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS

TABLE 1
Changes in the Distribution of Free Amino Acids and Amides in the Cotyledons of Acacia iteaphylla Seedlings

Age of scedling (days)

Compound 3 4 8 14
S-carboxycthylcysteine ++++ ++++ ++++ +++
S-carboxyisopropyleysteine +++ ++44 ++4+4+ 45 te fe
Pipecolic acid +++ ++4++ +44 de Le
Albizziine + asparagine pal ++ Rae soup a2
Glutamine ++ ++ See eet A tt
Aspartate tr ++ tr fe
Glutamate +++ +++ fe fe fe
Serine ++ + + ++ 4 ae
Glycine ++ + + +
Alanine ++ +++ ++ tr
Valine + tr tr tr
Leucine + Isoleucine + tr tr tr
Lysine tr tr tr tr
Arginine ++ + tr n.d.

tr, trace; n.d., not detected.
albizziine predominant;
asparagine predominant — sec text.

TABLE 2
Changes in the Distribution of Free Amino Acids and Amides in the Roots and Hypocotyl of Acacia iteaphylla Seedlings

Roots Hypocotyl
Compound 8-day 14 day 8-day 14-day
S-carboxycthylcysteine n.d. + ++ +++
S-carboxyisopropylcysteine n.d. n.d. n.d. ++
Pipecolic acid +++ ++ ++4++4+ +
Asparagine +++ ++ ++4++4+ +++
Glutamine ++ n.d. ++ n.d.
Aspartate + + ++ +
Glutamate + n.d. fp ae tr
Serine + ++ ++ +
Glycine tr ++ + n.d.
Alanine ++ + + +++ tr
Valine ++ n.d. + n.d.
Leucine + Isoleucine tr n.d. + n.d.
Threonine n.d. n.d. tr n.d.

tr, trace; n.d., not detected.

maintained, but the contents of pipecolic acid and alanine declined sharply (Table 2). In
the roots of the 14-day-old seedlings, some S-carboxyethylcysteine was present, but S-
carboxyisopropylcysteine was not detected.

In the balance of the shoots from these seedlings (mainly the primary leaf at 14
days), serine and S-carboxyethylcysteine were most prominent. Pipecolic acid was also
detected, along with aspartate, glycine, alanine, glutamate and glutamine, but
asparagine, albizziine and S-carboxyisopropylcysteine were not detected.

PROC. LINN. SOC. N.S.W.,, 111 (1), 1989

ANITA B. BRADKE AND DAVID R. MURRAY 41

DISCUSSION

The net increase in free amino nitrogen in the seedling between 1 and 8 days was
2.55umol (Fig. 1), which is within the maximum amount that could be met from limited
breakdown of proteins in the cotyledons (Ashcroft and Murray, 1979). Our observations
are consistent with the transport of selected forms of free amino nitrogen from the
cotyledons to the hypocotyl, the root, and later the shoot, with the content of free amino
nitrogen maintained close to 2umol per cotyledon by proteolysis until after 8 days (Fig.
1). It must be noted that it is not possible for any uptake of exogenous nitrogen sources to
have occurred through the roots; the axis-derived organs are gaining nitrogen entirely at
the expense of the cotyledons.

Since the only way that the cotyledons can export nitrogenous solutes is through the
phloem (Guardiola and Sutcliffe, 1972; Murray, 1984), an important question raised is
whether the different organs of the axis are selective in the nitrogenous solutes they
import, and if so, whether restrictions apply to phloem loading or unloading of in-
dividual compounds. In seedlings of jack bean (Canavalia ensiformis |L.] DC), Rosenthal
and Rhodes (1984) have demonstrated that the non-protein amino acid L-canavanine is
transported only to the above-ground parts of the shoot — none is translocated to the
radicle.

Of all the non-protein amino acids potentially available from the cotyledons, only
pipecolic acid appeared to be imported by very young roots of Acacza iteaphylla seedlings.
Pipecolic acid and asparagine were abundant early in both the roots and the hypocoty];
quantitatively, they are the most important forms of nitrogen initially transferred from
the cotyledons to the axis (Table 2). Outside the cotyledons, S-carboxyisopropylcysteine
was detected only in the hypocotyl, relatively late in development (Table 2), whereas
transport of the other derivative of cysteine was less restricted. S-carboxyethylcysteine
became one of the more prominent nitrogenous solutes of the hypocotyl, and appeared
also in the roots (late) and the primary leaf. The serine present in the primary leaf has
probably accumulated from at least two biosynthetic pathways operating in the leaf itself
(Murray, 1986a).

Arginine was not transported from the cotyledons (Table 2), but converted to other
nitrogenous solutes in situ (Table 1). Urease was detected immunochemically in
cotyledon extracts (for methods see Murray and Knox, 1977), which is consistent with
this interpretation.

Finally, what is the role of albizziine (L-a-amino-$-ureidopropionic acid), a com-
pound whose synthesis in Acacia cotyledons is known to be confined to seed development
(Seneviratne and Fowden, 1968)? In our chromatographic systems, albizziine and
asparagine could not be resolved (Fig. 2). This is in agreement with data on the mobility
of albizziine reported by Cooper and Meister (1973). However, the colour of the reaction
product with ninhydrin produced by authentic albizziine after paper chromatography
was always mauve, whereas that produced by asparagine was always brown. On
chromatograms where authentic albizziine and asparagine were loaded together, the
final colour reflected the abundance of each compound; a brown colour consistently
indicated the predominance of asparagine.

Initially albizziine is the prominent component of the combined albizziine-
asparagine area from cotyledon extracts, but with increasing time following germi-
nation, asparagine predominates. When detected on chromatograms of Acacia root and
hypocotyl extracts, the joint region was always brown, and if the unstained area was
eluted from replicate chromatograms and hydrolysed with HCl, a ninhydrin positive
product with the same mobility as aspartate was recovered. We have therefore con-
cluded that asparagine is a major nitrogenous solute translocated to the roots and

PROC. LINN. SOC. N.S.W., 111 (1), 1989

42 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS

hypocotyl (Table 2). This is in agreement with a transport function for newly synthe-
sized asparagine determined for cotyledons of light-grown pea (Melcher, 1983) and
peanut seedlings (Peoples et al., 1986).

In Acacia cotyledons albizziine, like arginine, is converted to translocated forms of
nitrogen, predominantly asparagine. It remains to be determined exactly how albizziine
is metabolized, and whether, while present in cotyledons, it has any deterrent function
against larvae of Australian insects.

ACKNOWLEDGEMENTS

The financial support of the Joyce W. Vickery Scientific Research Fund (Linnean
Society of N.S.W.) is gratefully acknowledged. We also thank Dr C. S. Evans and Prof.
E. A. Bell for helpful information and advice.

References

ASHCROEFT, W. J., and Murray, D. R., 1979. — The dual functions of the cotyledons of Acacia iteaphylla F.
Mucll. (Mimosoideac). Aust. J. Bot. 27: 343-352.

Cooper, A. J. L., and MEISTER, A., 1973. — Action of liver glutamine transaminase and L-amino acid
oxidase on several glutamine analogs. J. Biol. Chem. 248: 8499-8505.

EVANS, C. S., QURESHI, M. Y., and BELL, E. A., 1977. — Free amino acids in the seeds of Acacia spccics.
Phytochem. 16: 565-570.

GUARDIOLA, J. L., and SUTCLIFFE, J. F., 1972. — Transport of materials from the cotyledons during germi-
nation of seeds of the garden pea (Pisum sativum L.). J. Exp. Bot. 23: 322-337.

KRISHNA, T. G., and MuRRAY, D. R., 1988. — Effects of cycloheximide and actinomycin D on glycosidase
activities in the cotyledons of legume seeds following imbibition. J. Plant Physiol. 132: 745-749.
MELCHER, I. M., 1983. — Changes in nitrogen-containing compounds of the garden pea (Pisum sativum L.)

during germination. Z. Pflanzenphysiol. 112: 95-102.
Murray, D. R., 1981. — Functions of the cotyledons in Acacia. Aust. Plants 11: 65-66.

——.,, 1983. — Changes in free amino acid and amide composition during fruit and seed development of
garden pea, Pisum sativum L. New Phytol. 93: 33-41.
——, 1984. — Axis-cotyledon relationships during reserve mobilization. Jn MurRRay, D. R., (ed.), Seed

Physiology. Volume 2. Germination and Reserve Mobilization: 247-280. Sydney: Academic Press.
—, 1986a. — Amino acid and amide metabolism in the hulls and seeds of developing fruits of garden pea,
Pisum sativum L. 1V. Alanine. New Phytol. 104: 395-406.
——.,, 1986b. — Sced dispersal by water. Jn MuRRAY, D. R., (ed.), Seed Dispersal: 49-85. Sydney: Academic
Press.
, and KENNEDY, I. R., 1980. — Changes in activitics of enzymes of nitrogen metabolism in sccdcoats
and cotyledons during embryo development in pea seeds. Plant Physiol. 66: 782-786.
, and Knox, R. B., 1977. — Immunofluorescent localization of urease in the cotyledons of jack bean,
Canavalia ensiformis. J. Cell Sci. 26: 9-18.
—-, and McGrr, C. M., 1986. — Sced protein content of Australian species of Acacia. Proc. Linn. Soc.
N.S.W. 108: 187-190.
——., GIOVANELLI, J., and SMILLIE, R. M., 1971. — Photometabolism of glycolate by Euglena gracilis. Aust. J.
Bool. Sci. 24; 23-33.
PEOPLES, M. B., PATE, J. S., ATKINS, C. A., and BERGERSEN, F. J., 1986. — Nitrogen nutrition and xylem sap
composition of peanut (Arachis hypogaea L. cv. Virginia Bunch). Plant Physiol. 82: 946-951.
PETRIE, J. M., 1908. — The role of nitrogen and its compounds in plant metabolism. Part 11. The non-protein
nitrogen in seeds. Proc. Linn. Soc. N.S.W. 33: 835-844.
——,, 1911. — The role of nitrogen in plant metabolism. Part 111. The distribution of nitrogen in Acacza seeds.
Proc. Linn. Soc. N.S.W. 36: 97-126.
ROSENTHAL, G. A., and RHODES, D., 1984. — L-Canavanine transport and utilization in developing jack
bean, Canavalia ensiformis (L.) DC [Leguminosae]. Plant Physiol. 76: 541-544.
SCHNARRENBERGER, C., OESER, A., and TOLBERT, N. E., 1972. — Isolation of protein bodies on sucrose
gradients. Planta 104: 185-194.
SENEVIRATNE, A. S., and FOWDEN, L., 1968. — Diamino acid metabolism in plants with special reference to
a,@-diaminopropionic acid. Phytochem. 7: 1047-1056.
SMITH, D. L., 1981. — Cotyledons of the Leguminosae. Jn POLHILL, R. M., and RAVEN, P. H., (eds), Advances
in Legume Systematics. Part 2: 927-940. Kew: Royal Botanic Gardens.

PROC. LINN. SOC. N.S.W., 111 (1), 1989

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 111
NUMBER 2

John Vaughan Thompson, F.L.S.

HUGH CAMPBELL
(Communicated by T. G. VALLANCE)

CAMPBELL, HUGH. John Vaughan Thompson, F.L.S. Proc. Linn. Soc. N.S.W. 111 (2),
1989: 45-64.

John Vaughan Thompson (1779-1847) was a surgeon in the British Army Medical
Service who made his name as a ‘gentleman naturalist. He became a Fellow of the
Linnean Society in 1810 in recognition of his work in botany, but his great contribution
to science was his discovery of metamorphosis in the Crustacea: a discovery which was
received with disbelief and hostility in the 1830s, and not fully accepted until after
Thompson died. He spent the last eleven years of his life in New South Wales, eight of
them in charge of Convict and Military Medical Services. He failed as an adminis-
trator, and most Australian historians have not only blamed him unduly for his failure
but have also neglected the scientific achievements which more than compensate for it.
His life is an interesting study of the difficulties faced by an amateur scientist working
alone without the support of personal wealth or powerful friends.

N. H. Campbell, 19 Brushy Creek Road, Lenah Valley, Australia 7008; manuscript received
19 July 1988, accepted for publication 23 November 1988.

INTRODUCTION — GIVING J. V., THOMPSON HIs DUE

John Vaughan Thompson (1779-1847) distinguished himself as a zoologist and
botanist while quietly earning his living as an army surgeon in England, the West
Indies, Mauritius, Madagascar, Ireland and Australia. He has been treated generously
by his recent English biographers, who refer to his achievements before he went to Aus-
tralia. Australians have been less kind to him: if they have noticed him at all, it is only in
passing. For instance, although he spent his last eleven years in New South Wales, eight
of them in charge of the convict and military medical services in the colony, the ADB
contains no entry about him in his own right: he is mentioned only as a sort of béte nozre in
other men’s lives.

It is true that he failed as a colonial administrator, and only natural that local
historians should have reported accordingly. But most did not inquire further, and so
overlooked both his zealous service during forty-five years as an army surgeon and his
wide-ranging work in natural history, especially his discoveries about marine in-
vertebrates. What is more, the writer believes they made their unflattering picture even
worse by blaming him too much for his difficulties in New South Wales. As a result,
biographers sympathetic to Thompson have usually thought it best to dismiss his years
there in a single sentence and concentrate on the rest of his life. This paper is an attempt
to repair the large gaps and unfair biases in what has hitherto been written about him.

There are other reasons, too, for the discrepancies. First, Thompson was inconsis-
tent in himself: illness and bitterness changed him during the course of his service in
New South Wales. Second, earlier biographers do not mention the Colonial Office files,
reported for the first time in this paper, which contain a number of details of his personal
life. Hence, they do not explain some of his actions. Third, several of his discoveries in
natural history soon became such common knowledge that his later biographers forgot
his part in discovering them.

Wheeler (1970) pointed out that most of Thompson's important achievements were
recognized too late for him to enjoy the praise: he had that sort of luck all his life. By his
own account, fortune first failed him in 1778, the year before he was born, when his
father lost all his property in North America. In one part of a ‘memorial’ seeking a grant

PROC. LINN. SOC. N.S.W., 111 (2), 1989

46 JOHN VAUGHAN THOMPSON, F.L:S.

of land in New South Wales, Thompson (1835) explained the setback: ‘Memorialist’s
father . . . lost to his family by attachment to the Loyalist cause and the unfortunate
result of the American War, a grant of land to the amount of 4,000 acres situated on the
Mohawk River beyond Albany, . . . together with a good deal of other property, houses,
&c at New York and Long Island, and a considerable stock of firing provided for the
army under Sir John Vaughan, for which he was contractor — the above grants being
given as the reward of services as an officer of the 96th Regiment at the taking of Pt
Havanna [sic] &c was totally lost by the precipitation with which he was obliged to fly
when the King’s troops took their final departure from Sandy Hook’ .*

It was typical of Thompson to expect the government to compensate him for the
loss his Loyalist father had suffered in the American War of Independence more than
fifty years earlier — a loss he had probably heard of many times as a boy. But, as we shall
see, to the end of his life, he remained surprisingly naive in political matters.

On 19 November 1779, a year after his father fled from Sandy Hook, John Vaughan
Thompson was born at Berwick-on-Iweed. His mother was Jane Hall (International
Genealogical Index, 1981).

Thompson grew up there, and gave that address as late as 1817. But he left home
much earlier: the son of a dispossessed Loyalist army officer had to make his own way in
the world. He studied medicine at the University of Edinburgh in the sessions of 1797-8
and 1798-9, enrolling in anatomy, surgery, obstetrics, chemistry and botany (Edinb.
Univ., 1797-8); and on 3 March 1799, soon after his nineteenth birthday, he was
appointed Assistant Surgeon to the Prince of Wales Fencibles. Without taking his
degree, he accompanied the regiment to Gibraltar in December 1799.

THE WEST INDIES, 1800-1809

Appointment as Army Surgeon

On 3 July 1800 he became Assistant Surgeon of the 37th Foot, and embarked with
them for the West Indies and Guiana to take part in the war against the French and
Dutch: records show that he was engaged in the taking of Berbice and Demerara in
Guiana. On 25 June 1803 he was promoted to Surgeon in the regiment. During his six
years residence in Trinidad, with occasional visits to Grenada and St Vincent, he found
time, while carrying out his military duties, to study the natural history of subjects as
diverse as land crabs and ginger.

There is evidence that he paid at least one visit to England during his service in the
West Indies. When he asked Lord Seaforth, F.L.S. and recently retired Governor of
Barbados, to present his paper about Kaempferia to the Linnean Society, he headed his
covering letter ‘London, April 7th, 1807’ (Linn. Soc., 1807). He took advantage of being
in England that year to engage in other literary activity: he published his ‘Catalogue of
Plants in the vicinity of Berwick-on- Tweed’ (a small octavo of 132 pages) (Thompson,
1807a), and had papers on Kaempferra (Thompson, 1807b) and Piper (Thompson, 1808)
read to the Linnean Society.

Early Interest in Botany

Britten (1912), reviewing his first publication, wrote: “Thompson must early have
acquired considerable proficiency in botany. His Catalogue of Plants ... Berwick upon
Tweed was prepared before he left England — i.e. at or before the age of twenty — and

* The Encyclopedia Americana (1983) explains the reference to the capture of Havana in 1762; the DNB (1900a)
entry on Sir John Vaughan gives an account of his service on the British side in the American War of Indepen-
dence and his appointment as Governor of Berwick-on-Iweed, where Thompson’s father appears to have
joined him, a possible connnection by a marriage in 1765 being suggested by the International Genealogical Index
(1981); and the EB gives an account of the British flight from Sandy Hook in 1778.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 47

shows a very complete knowledge of the plants of that region and of the literature of the
period: the pretty coloured group which adorns the title page and a plate (also coloured)
and dissections show him to have been a capable artist’.

In the same article, Britten points out that a paper on British birds has been
wrongly attributed to Thompson (DNB, 1900b) because of a false entry in Royal Society
(1871).

Thompson left the West Indies for good and returned to England in 1809. A year
earlier he had been made an Associate of the Linnean Society, and in November 1809,
four members signed the following certificate: ‘John Vaughan Thompson, Esq., Surgeon
of his Majesties [sic] 37th Regiment of Foot, & A.L.S., a Gentleman well versed in the
study of Botany, being desirous of becoming a Fellow of the Linnean Society of London,
we, the undersigned do from our personal knowledge beg leave to recommend him as
likely to become worthy of that honour (Linn. Soc., 1809).*

He was elected F.L.S. on 6 February 1810. Stebbing (1910), writing on the centenary
of the occasion, described his election as ‘prophetic insight’, recognizing that
Thompson's most important work was still to come.

On 3 March 1812, two of his papers were read to the Society — one about an
unusual Pouched Rat he had observed in Trinidad (Thompson, 1813) and the other,
reported by Wheeler (1970), on his observations of the genus Myrti there.

SERVICE IN MAURITIUS AND MADAGASCAR, 1812-1816

Shortly afterwards, he was posted to Mauritius and Madagascar. On Christmas
Day, 1812, he was appointed staff surgeon, though his official duties in the Mascarene
Islands are not now known. It has been said that in two consecutive years he worked on
‘Introducing vaccine innoculation to Madagascar, and that for some time he was
officially styled ‘Government Agent for Madagascar’. In his 1835 letter seeking a land
grant (Thompson, 1835), he himself refers to his appointment instructions and his jour-
nal ‘sent to Governor Sir R. T. Farquhar and now, he presumes, in the Colonial Office’.
Neither the instructions nor the journal have been found.

He claims (Thompson, 1835) that: ‘amongst the important accessions to the culture
of the Mauritius, [he] added 8 varieties of that estimable food, the plaintain, 10 valuable
varieties of banana, 12 varieties of sugar cane, most of them vastly superior to those in
common cultivation, 9 varieties of tobacco, 7 varieties of the indigo plant and 11 varieties
of ordinary and upland rice’.

He also writes of: ‘His efficient service as Civil Agent at Madagascar for the years
1814-15 a. During the period of his Agency he put a total stop to the traffic in slaves from
those parts under his control. b. Repressed several efforts of rival chiefs to disturb the
general tranquility [sic] by war. c. Introduced vaccine innoculation amongst the natives.
d. Completed a survey of the extensive harbours of Loquez without loss of a single life. e.
Paved the way for a friendly intercourse with the natives by a kind and generous treat-
ment diametrically opposed to that of the French, our predecessors. f. Introduced and
taught the cultivation of the potato and several other culinary vegetables’.

Vaughan (1953) refers to a 1902 work by Grandidier in which Thompson was
credited with introducing Albizzza lebbek (‘bois noir’) to the treeless plains of Madagascar,
having included a sack of seeds among his presents to the island’s king, Radama I. The
reference is further evidence that Thompson was engaged in some sort of diplomatic
service in Madagascar.

* Apparently wishing, on second thoughts, to show their wholehearted endorsement of an Associate, the
sponsors struck out ‘likely to become’.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

48 JOHN VAUGHAN THOMPSON, F.L:S.

Thompson says that eventually he was ‘attacked by the formidable remittent fever
of the country which obliged him to relinquish a very lucrative post to return to Europe
at great expense and finally to go on the half pay of his then rank of surgeon’.

In his informative ‘land grant letter, he also makes a particular point of mentioning
that he had ‘at the request of Governor Farquhar, drawn up and published at the
Mauritius a systematic catalogue of all the plants cultivated in the three Government
Gardens of Pamplemousses, Mon Plaisir and Reduit, which cost [him] much labour,
numerous journeys and a great deal of patient investigation’.

The title page of the Catalogue of Exotic Plants Introduced to Mauntius (Thompson?,
1816) does not show Thompson as the author. R. E. Vaughan (1953), however, based an
article entitled A Forgotten Work of John Vaughan Thompson on yet another letter, which
Thompson had written about the Catalogue in 1838. In that letter, he claimed to be the
author of the work, which had in fact been published anonymously in 1816, shortly after
he had left for England. He asserts that someone had suppressed the title page naming
him as author, which had been ready for the press when he sailed, and replaced it with
another. The 1838 letter was an appeal to the President of ‘La Société d’Histoire
Naturelle de Maurice’ asking the Society to credit Thompson with the authorship. In
the event, the President did nothing to advance Thompson's claim, partly because he
thought that the Catalogue, based as it was on the Linnaean ‘systeme sexuel, was out of date
by 1838.

Regardless of authorship, ‘this was the first work on the plants of the island to be
published locally, and is useful in listing the dates of introduction of many plants grown
in the Botanic Gardens and elsewhere. It shows Thompson’s interest in the importation
of useful plants, an interest he shared with many. other colonists of the time’ (Wheeler,
1970). In fact, it contains references to a number of plants introduced by Thompson
himself. Barnwell (1941) lists among these the honeysuckle, the jonquil, Guernsey and
other lilies, Madagascar arrowroot and tobacco, weeping willow, walnut, chestnut,
horse-chestnut and American lime; he also records that Thompson’s name is inscribed
on the Liénard Obelisk in the Pamplemousses Garden. But this first taste of having his
scientific efforts and their worth doubted probably influenced Thompson’s later decision
to publish his own work.

Britten (1912) reports that during this period he also ‘sent dried plants to Robert
Brown, which are in the National Herbarium’, later to become part of the British
Museum (Natural History).

Reviewing his service between 1812 and 1816, Thompson later complained of the
‘loss of property at the Madagascar and the Mauritius, being involved in the latter by the
burning of Port Louis* and the extensive bankrupcies [sic] which occurred shortly after
his departure for Europe on sick leave’, and of the ‘loss and expense entailed by the great
length to which the regnal|[?] of the Madagascar was extended’.

When Thompson left Mauritius, he was thirty-seven. In the year of his departure,
he suffered physically from the onset of malaria, financially from the fire at Port Louis,
and psychologically from lack of recognition of his efforts. And his hardship was com-
pounded because illness forced him to go on half pay for two years from 10 June 1817
(Johnston, 1917).

He had the consolation, however, of knowing that his paper on Mus anomalis, which
had been read before he left England, had been published by the Linnean Society in
1813 (Thompson, 1813). The paper, accompanied by a plate which illustrates his

*“A fire broke out in the commercial section of Port Louis [in 1815]. Efforts to extinguish it were unsuccessful
and all the principal shops and warchouses were destroyed. Seven hundred houses were burned down, and
millions of rupees of property lost. [Governor] Farquhar acted quickly and efficiently. People were clothed
and fed and no one died of hunger, although many were ruined financially . . .’(Mannick, 1979).

PROC. LINN. SOC. N.SW.,, 111 (2), 1989

HUGH CAMPBELL 49

meticulous drawing, contains a revealing statement of his views about scientific classifi-
cation: ‘But when we examine into nature with due attention, we find she delights to
mock the vain efforts of mortals to shackle and confine her within the bounds of generic
characters, which are found to run so into each other as to render all attempts at method
more or less imperfect. This animal must remain an anomaly in the family’.

His curiosity about natural history was not dampened. He had taken advantage of
his time in Mauritius to study ‘the famous extinct Mascarene birds’, although he waited
until 1829 to publish his Contributions towards the natural history of the Dodo . . . (Thompson,
1829). More immediately, as Wheeler (1970) records, on the way home he observed
south of Madagascar ‘a puzzling luminosity in the sea. He trailed a muslin hoop over the
stern of the ship and caught a profusion of small animals hitherto invisible in the water’.
Wheeler goes on to note that “Thompson has been credited with being the first to use a
plankton net, and there is little doubt that his use of it in late July or August 1816 was his
own idea entirely: but he was anticipated by John Cranch, who used a similar tow net on
[Captain Tuckey’s voyage to] the River Zaire (or Congo) in April 1816.

Back in Berwick-on-Iweed on half pay, Thompson returned to medical school. He
enrolled for the 1816-17 session at the University of Edinburgh in Clinical Medicine and
Materia Medica. He paid 4/6d for lectures in Materia Medica from Professor James
Home; among others on his page of the fees list were two Royal Navy surgeons and
another army surgeon (Edinb. Univ., 1816).

ARMY MEDICAL SERVICE AND MARINE STUDIES AT CORK, 1819-1835

The next record of Thompson’s movements appears when he was restored to full
pay (on exchange) on 25 May 1819 (Johnston, 1917). By that time he was in Cork, and
for the ensuing eleven years he remained on full pay in posts such as district medical
officer at Cork and surgeon of the Cork Recruiting District. There is evidence that in
1819 he gave a course of lectures at the Cork Institute on the science of Botany (Wheeler,
1977/5)

Presumably he married soon after his return to England in 1816. The advice of his
departure from Mauritius in June 1816 (Gazette de l’Isle Maurice, 1816) suggests that he
had no wife then. Previous biographers have said nothing about his private life and
family; Wheeler (1970), the latest, said, ‘It is not known whether he married’ But it is a
subject on which Thompson’s (1835) revealing letter throws some light. In it, he
describes himself as ‘having a young family of six children, four girls and two boys,
whose prospects have been materially affected’ by the reverses in North America and
Mauritius which he mentioned in the letter. Tantalizingly, the shipping record shows
that, when he sailed for Hobart Town in 1835, he was accompanied by his wife and four
children, the eldest sixteen (Colonial Office, 1836a): for some reason, two of the girls
(probably older than 16) did not go.

Thompson remained at Cork until 1835. During his years there he made those
discoveries of the life histories of the marine invertebrates in the Cove [Cobh] for which
he is chiefly remembered. He announced the principal ones in the following order
during the brief period between 1827 and 1830:

the discovery of Pentacrinus europaeus (Thompson, 1827);
t the discovery of metamorphosis in the development of most Crustacea (Thompson,
1828);
the classification and life history of barnacles (Thompson, 1830a); and
the discovery of the animals he called Polyzoa (Thompson, 1830b).
As the DNB (1900b) has it, ‘Our present conceptions of the structure of these forms, of

PROG. LINN. SOG. N.S.W., 111 (2), 1989

50 JOHN VAUGHAN THOMPSON, F.LS.

their zoological position, and of the metamorphoses which they undergo, date from
Thompson's papers.

His achievements resulted from his appreciation of the need for practical obser-
vation, his capacity for detecting and recording detail and, above all, his ability to
realize the implications of what he saw. They were all the more remarkable because he
worked alone, without the support of a university or other centre for scientific develop-
ment, or the collaboration, it seems, of even a single colleague. Hence, for want of
people to rally round him in his lifetime, his name is not well known today, even among
specialists in marine zoology. Although his discoveries about marine invertebrates
received a good deal of attention when he announced them, biologists soon began to
take them for granted: and if scientists subsequently wondered who made those dis-
coveries, they often attributed them to later workers.

Thompson announced his discoveries in a series of memoirs which he published at
his own expense. Although nowhere does he say so, it is likely that his experience with
his Catalogue of Exotic Plants in Mauritius made him wary of trusting his discoveries to
others until he had published them.

Pentacrinus europaeus

The DNB (1900b), in its reference to his first discovery, the Memoir on Pentacrinus
europaeus (Thompson, 1827), records that it: ‘announced the presence of a stalked crinoid
in the seas of the British Isles: until then crinoids (feather stars or sea-lilies) were known
only from the West Indies. Thompson also revealed that the crinoidea were really
‘radiata’; and (as shown more fully by a second paper in New Philosophical Transactions,
Edinburgh 1836) that this pentacrinus was really the young stage of the antedon, or feather
star’.

The discovery of this animal and Thompson’s conclusions about the stages in its life
history drew the attention of zoologists in France, Germany and elsewhere to his work,
and many of his succeeding papers were translated or abstracted into journals abroad.

Thompson announced his other principal discoveries in a series of five scarce
pamphlets, now reprinted in facsimile, which he published under the title of Zoological
Researches and Illustrations (Thompson, 1828-34).

In the ight of Thompson’s later appointment to Sydney, it is interesting to note that
a copy of the first Researches was sent to New South Wales, endorsed ‘W. S. McLeay,
Esq., with the author’s compliments. ‘He was also the donor of seeds of cotton to A.
McLeay in Sydney, as shown by the Seed Book’ (Fletcher, 1920). In fact, Thompson
corresponded regularly — and evidently was on familiar terms — with Alexander
Macleay, the Treasurer of the Linnean Society of London from 1798 until 1825, and
thereafter Colonial Secretary of New South Wales (Wheeler, 1970). Macleay, of course,
was one of the four who had nominated him for F.L.S. in 1809.

Metamorphosis of Crustacea

Thompson published the first issue of his Researches at Cork in September 1828,
price 3s.6d. ‘It announced what is probably his most important contribution to zoology:
the discovery that certain planktonic forms of crustacean, then known by the generic
name Zoea, undergo changes of form (metamorphoses) until they become recognizable
as the young of the European edible crab (Cancer pagurus) (Wheeler, 1970).

The full title of the first memoir was: On the Metamorphoses of the Crustacea, and on Zoea,
exposing their singular structure and demonstrating that they are not, as has been supposed, a peculiar
genus, but the Larva of Crustacea!! (Thompson, 1828). Wheeler (1975) points out that ‘the
two exclamation marks as well as his text showed that he appreciated the revolutionary

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 51

nature of his discovery, but it is doubtful whether he could have anticipated the furore
that his important discovery was to invoke among the zoological establishment’.

Wheeler (1975) goes on to explain how Thompson’s discovery challenged one of the
basic tenets of contemporary systematic zoology: “The essence of the controversy lay in
the distinction made by systematists between the classes /nsecta and Crustacea. The
insects, as was easy to observe, went through a series of metamorphoses to emerge as an
imago, or perfect insect. The crustaceans, less easy to observe, were not known to
develop through metamorphosis; consequently they were assumed not to do so. This
difference was one of the principal characters distinguishing the classes’

Wheeler’s summary of Thompson’s memoir is simple and elegant: “Thompson’s
demolition of the character was based on practical observation. In 1816, while on the
return journey from Mauritius, he had . . . captured small planktonic animals, which
resembled species of the crustacean genus Zoea, described by earlier naturalists from
mid-ocean. In the spring of 1822, Thompson “to his great surprise” found a considerable
abundance of Zoeas in the harbour of Cove [Cobh], and in 1823 he succeeded in keeping
a large specimen in fresh sea water between 14 May and 15 June, when it died in the
process of casting its skin. He noted that the form of the Zoea was totally changed and
that the limbs that had disengaged from the skin resembled those of the decapod crusta-
ceans (crabs, lobsters, shrimps, etc.) in having four long walking legs and an anterior
pair armed with pincers. The final confirmation was obtained four years later, in June
1827, when Thompson succeeded in hatching the eggs of the edible crab (Cancer pagurus)
to find zoea larvae. From these observations he asserted that the decapod crustaceans
generally undergo metamorphosis during their development. Not only did his
announcement render invalid ong of the major distinctions between the Crustacea and
the Insecta, it accounted for the anomalous Zoea specimens of earlier naturalists, and it
also explained previously unaccountable phenomena such as the migration to the sea of
land crabs which he had observed in the West Indies’.

In fact, Thompson used his observations of land crabs to support his theory, point-
ing out that even such land dwellers must lay their eggs in the sea and give their young a
perilous life there, so that they may undergo metamorphosis before going ashore in their
adult form.

He points out, too, that fifty years earlier, in a work published at Haarlem, Slabber
(1778) claimed that he had observed metamorphosis in Zoea. Thompson swiftly disposes
of that claim: “The metamorphosis, however, which this observer thought he witnessed is
of so different a description that we must either suppose him to have fallen into some
error or else there may be Crustacea which pass through some other forms’

He concludes, ‘from much experience’, that Slabber ‘lost his Zoeas in changing the
sea water, and that the new form came from the added portion’.

Nevertheless, in his own first memoir, Thompson, too, had ‘failed to prove the
complete metamorphic cycle because his zoea died in the process of change; it was only
by comparing them [sic] with ova from a berried female crab that he was able to deduce
the relationship (Wheeler, 1970). What is more, his announcement was accompanied by
a second memoir ‘On the genus Mysis (Thompson, 1828), in which he showed that the
mysidacean crustaceans hatch in a form very similar to that of the adult and undergo
only slight metamorphosis, thus tending to contradict his assertions about metamor-
phosis as a characteristic of most crustaceans. And both memoirs appeared only months
before the German scientist Rathke, unaware of Thompson’s announcements, pub-
lished his own observations on the development of the crayfish (Potamobius, formerly
Astacus), now known to be one of the few exceptions to the general rule propounded by
Thompson. Rathke’s work demonstrated that the young crayfish hatch at a late stage of
development and do not undergo metamorphosis.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

52 JOHN VAUGHAN THOMPSON, F.L:S.

One might ask why Thompson was in such a hurry to publish when his conclusion
was not entirely proved and was supported by only a single observation. The answer lies
in the comment made by Stebbing in relation to another of Thompson’s discoveries, the
Polyzoa, where an observation made in 1823 was not published for seven years, and so
was not credited to him. ‘As we all know, recognition of our discoveries has to date, not
from the time when they were made, but from the time when they were published’
(Stebbing, 1910). Thompson knew that others were studying marine invertebrates, and
did not want to risk being beaten. As it happened, he was alone in detecting metamor-
phosis, and could have afforded to wait.

But given Thompson’s reliance on deduction in his first memoir, it is not surprising
that many established zoologists treated his claim of metamorphosis in the Crustacea
with ‘distinct and often derisory doubt. They resented the overturning of the taxonomy
in use at the time, as expounded by Leach and Cuvier. Nevertheless, Thompson con-
tinued to announce fresh examples to prove his hypothesis. By the end of the decade,
Rathke and others had published admissions that their scepticism had been misplaced
and that Thompson was correct (Wheeler, 1975).

In the middle of 1830, Thompson’s military career blossomed briefly. On 22 July he
was promoted to the rank of Deputy Inspector General of Hospitals (the next step on the
promotion ladder for surgeons) as a reward for his long and zealous service. The pro-
motion promised him an income sufficient both to support his family in the style he
thought fitting and to continue publishing the results of his research. Then, as suddenly
as his fortunes rose, they fell (Johnston, 1917). On 9th December 1830 he was retired to
half pay, along with 13 other officers of his rank who experienced the same meteoric rise
and fall. At age 51, with a wife and six children (under fifteen?), he saw his income drop
from 30 shillings a day to 17. Not only did the reduction threaten the prospects of his
family, but it also made it impossible for him to go on publishing his Zoological Researches
without help. Some might think he should have been able to bring up a family on £300 a
year. Thompson did not.

Consequently, the proceedings of a meeting of the Zoological Society of London a
month later, on 11 January 1831, (Zool. Soc., 1831), show that it received ‘an Address by
Mr J. V. Thompson, Esq., “Io the members of the Zoological Society, and the Zoologists
of the United Kingdom in general”, soliciting such support, by subscription, as may
enable him to continue, without further loss, his Zoological Researches and Illustrations. This
address is printed, together with a list of the subjects of some of the preceding memotrs,
on the cover of the fourth number of the Researches, which was at the same time laid on
the table’. Stebbing, writing eighty years later, referred to ‘a pathetic appeal to the scien-
tific world to furnish him with 150 subscribers, as his private income would no longer
bear the sacrifice till then entailed by the publication of his researches’ (Stebbing, 1910).

Thompson’s financial setback did not prevent him from announcing further
evidence of metamorphosis in the Crustacea, in response to a complaint from Vigors
(1830) in the Zoological Journal that his conclusions had been too sweeping. His letter to
the Secretary of the Zoological Society dated ‘Cork, Dec. 16, 1830 is reported thus
(Zool. Soc., 1830: internal quotations are from Thompson): ‘In it, Mr Thompson urges,
in support of the universality of a metamorphosis among the C7ustacea, that he has ascer-
tained the newly hatched animal to be a Zoea in eight genera of the Brachyura, viz. Cancer,
Carcinus, Portunus, Eriphia, Gecarcinua, Thelphusa?, Pinnotheres, and Inachus; and in seven
Macrourous genera, viz. Pagurus, . . . and Astacus. “These embrace all our most familiar
native genera of the Decapoda”. The Lobster, or Astacus Marinus, Mr Thompson states,
“does actually undergo a metamorphosis, but less in degree than in any of the other
enumerated genera, in its first stage being what I should call a modified Zoea . . .; in
short, such an animal as would never be considered what it really is, was it not obtained

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 35)

by hatching the spawn of the Lobster”. In the other indigenous species of Astacus, the
River Crawfish, it would appear from the excellent treatise of M. Rathke on the de-
velopement [sic] of its eggs, that the young are hatched in a form according with that of
the fully grown animal. Mr Thompson, however, suspects that some source of error may
exist in these observations. “If it should be found otherwise, it can only be regarded as
one solitary exception to the generality of metamorphoses, and will render it necessary
to consider these two animals for the future as the types of two distinct genera”. . .’

This letter is an interesting demonstration of Thompson’s confidence in the
accuracy of his observations: he did not hesitate to question the opinions and obser-
vations of established and respected naturalists when they conflicted with his own.

The Cirripedes

Thompson’s third important achievement in marine biology was his discovery that
cirripeds are Crustacea: in the system proposed by Cuvier, they had been designated as a
class of the Mollusca. He published his discovery in the third number of Zoological
Researches, Memoir iv: On the Cirripedes or Barnacles; demonstrating their deceptive character, the
extraordinary Metamorphosis they undergo, and the Class of Animals to which they indisputably
belong (Thompson, 1830a).

Wheeler (1975: internal quotation is from Thompson and Darwin) explains the
significance of the discovery thus: ‘Again this presented a revolutionary concept as far as
the higher classification of barnacles was concerned. Among the barnacles two major
groups are well known, the goose or ship’s barnacles, which look somewhat like small,
light coloured mussels on a stalk, and the acorn barnacles, which are so abundant on
rock and timber on shore. Earlier naturalists had been much exercised about the true
relationships of these animals.

“Thompson stumbled on the correct solution to their relationship by, as he put it,
“the result of chance rather than of design and industry”. In 1823, crossing on the ferry at
Passage Cove, he trailed his small muslin net behind the boat to catch a quantity of
marine zooplankton. Among them were translucent elliptical animals a tenth of an inch
in length, evidently crustaceans but of a form undescribed by earlier naturalists. On 1
May 1826 he collected further specimens, and in the light of his knowledge that larval
decapod crustaceans were entirely dissimilar to the adults kept them in order to observe
any metamorphosis which might take place. He had the satisfaction on 8 May of seeing
some of these elliptical animals metamorphose, and found them firmly attached to the
bottom of the glass container, perfectly recognizable as young acorn barnacles. On 10
May another of these larvae was seen to cast its outer skin and settle to the bottom as a
barnacle. In 1835 Thompson described the larvae of the goose barnacle, which he found
on a ship’s bottom in 1830.

‘Again, though Thompson had firmly placed the barnacles within the Crustacea
and showed that they too enjoyed a planktonic larval stage before settling, established
zoologists were slow to accept his findings. However, Darwin (1851) in his monograph on
the group recognized Thompson’s contribution as a “capital discovery” .’

Of course, Thompson would not have ‘stumbled on the correct solution had he not
been looking with the eyes of one who knew that metamorphoses occurred in Crustacea.

The Royal Society’s Catalogue of Scientific Papers 1800-1863 (Royal Society, 1871)
records some of the reprints of Thompson’s papers which had appeared by 1831 in
English and French journals.

An illustration of the way Thompson’s name cropped up incidentally in scientific
discussions in the early 1830s comes from the proceedings of the Zoological Society on
11 January 1831 (Zool. Soc., 1831). A question was raised at this meeting about the cause
of phosphorescence in sea water. ‘It was remarked that Commerson and others have

PROG. LINN. SOG. N.S.W., 111 (2), 1989

54 JOHN VAUGHAN THOMPSON, F.L:S.

attributed the phenomenon described to putrefaction of animal matters, . . . Sir Joseph
Banks, Dr Macartney, and others, on the contrary, have referred it to the presence of
marine animals, principally Crustacea; and the existence of such, as the cause of this
appearance, has been recently insisted on by Mr J. V. Thompson,’

Polyzoa

The last of Thompson’s four important discoveries was that of Polyzoa, a new Animal
discovered as an Inhabitant of some zoophytes (Thompson, 1830b). He showed that the term
zoophytes had been used to cover a mixture of animals superficially alike but essentially
different in structure. Relying on Wheeler again: ‘[ Polyzoa| had been formerly included
as part of a heterogeneous collection of enigmatic invertebrates, the so-called zoophytes;
but he showed that they were distinct from the colonial hydroids and the ascidians, with
which they had been sometimes confused. The term Polyzoa received considerable usage,
especially in Great Britain, but it was eventually dropped in favour of Bryozoa, which had
been proposed almost contemporaneously’ (Wheeler, 1970; supported by Stebbing,
1910).

Because of Thompson’s preoccupation with natural history, it is not surprising to
find that he left only one paper on medical science. Although he was a prolific writer and
despite more than forty years in the Army Medical Department, his only recorded pub-
lication in the field is an 1832 pamphlet entitled The Pestelential Cholera Ummasked . . .”
Wheeler (1970) describes it as ‘a work devoted to diagnosis and treatment of cholera, but
exhibiting little understanding of the causative factors involved in the disease’. Perhaps
the critic expected too much. At the time, no one else understood such matters, either. In
fact, the pamphlet conveyed the prevailing wisdom about the disease and it was topical:
cholera had been spreading westward from Asia during the 1820s, and the first major
outbreak in the British Isles occurred in 1832.

Wheeler (1975) reports in passing that in 1833 Thompson's collection of in-
vertebrates was purchased by the Royal College of Surgeons, Dublin. Unfortunately, the
collection has not survived. In the light of his financial setback at the end of 1830, one
wonders what domestic crisis forced Thompson to sell it.

SCIENTIFIC DISPUTES AND FINANCIAL DIFFICULTIES

Meanwhile, Thompson’s conflict with the zoological establishment in London
became more and more acrimonious as his new concepts threatened to overturn
fundamental systematics. As Wheeler (1970) says, ‘His correspondence and published
writings of the time suggest a man impatient with the conservatism of his opponents,
and eventually embittered by their opposition . . .’ Elsewhere, Wheeler (1975) says: ‘His
tendency to write in a forthright style cannot have endeared him to his opponents’.

Meanwhile, too, Thompson’s money problems became more and more embarrass-
ing. On 8 May 1835 he informed the Treasurer of the Linnean Society that he was
‘unable to send the large amount due’ and ‘craved the indulgence . . . in the hands of the
Committee of remitting the payments in special cases of disability. He pointed out that
he had not been attending meetings or receiving copies of the Society’s ‘valuable Trans-
actions’ while his fees (at the time £3 per annum) were in arrears. He explained that he
was claiming the indulgence ‘. . . on account of the pecuniary loss sustained by the pub-
lication of my Zoological Researches and the great expense incurred by the prosecution of
those discoveries by being obliged to keep up an establishment on the sea-side during a
great part of the period, & the disbursements consequent on boat hire travelling back
and forward, &c (Linn. Soc., 1835).

According to its minutes for 22 May 1835, Council ‘resolved that the request be

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 55

acceded to on account of his high merit as a naturalist and the papers he had supplied for
the Transactions of the Society.

In the middle of dealing with both his financial difficulties and a bitter scientific
controversy, Thompson was removed from the scene. Colonial authorities in London
had for some time been anxious to reduce the cost of the separated convict medical ser-
vices and military medical services in New South Wales and Van Diemens Land. Early
in 1835, the Government decided to combine both services in each colony under the
supervision of senior Army surgeons (Colonial Office, 1834). Accordingly, in June 1835,
John Vaughan Thompson was offered an appointment as Deputy Inspector-General of
Hospitals in New South Wales (Colonial Office, 1835a). Distressed as he was, he could
not refuse the offer, since it meant he would be restored to full pay from 7 August 1835
(Johnston, 1917).

Only money could have taken him so far abroad; and it could not have done so at a
worse time, because scepticism about his Zoological Researches was at its height. In the very
week that Thompson was offered his new appointment, his most implacable opponent,
J. O. Westwood, F.L.S., secretary of the Entomological Society and through that office
committed to retaining the old taxonomy, attacked Thompson's work by presenting a
long paper to the Royal Society On the supposed existence of Metamorphoses in the Crustacea.
Westwood (1835) used special cases and the opinions of a number of respected authori-
ties to deny outright the evidence that Thompson presented. He was quite wrong, but
the error did not make his criticism any less credible to many of his listeners and readers,
who shared his misconceptions. Patronizingly, he pointed out that ‘the accuracy of
[Thompson's] beautiful figures deserves the highest praise’ and summed up by saying,
‘Although disagreeing with Mr Thompson in respect to his theory, I have already stated
that his figures are very faithful delineations of nature’. In all else, he set out to discredit
Thompson.

Perhaps that paper spurred others, some of them also severe critics, to take
Thompson seriously and make their own observations. Nevertheless, it took some time
to verify Thompson's work, and it was not until the end of the decade that impartial zool-
ogists repeated his observations and admitted that he was right (Wheeler, 1975).

Not that Thompson himself had slackened in his efforts to substantiate his dis-
coveries. He had been continuing his research and his writing, and a number of his
papers appeared in various journals in 1835 and 1836 (Royal Society, 1871). Some of
them appeared even in the heart of the enemy’s territory, the Entomological Magazine
(Royal Society, 1871).

DEPUTY INSPECTOR-GENERAL OF HOSPITALS,
NEW SOUTH WALES, 1836-1844

Surgeon-Superintendent of the ‘Boadicea’

In June 1835, however, Thompson had the more pressing problem of arranging
passage to New South Wales for himself, his wife and four children. In his fifty-sixth year
he was about to begin a new career in a new land, and turn his back on the researches
which had occupied him for the past 20 years.

By the beginning of August 1835 he had settled most of his affairs in Ireland and
returned to London. He left unpaid a seemingly trivial account for goods to the value of
£6.10.7% with Mrs Dwyer of 89 Grand Parade, Cork, which was to haunt him for years
(Colonial Office, 1841a). On 7 August 1835, the day he returned to full pay, he wrote a
memorial to the Secretary of State for the Colonies, seeking a grant of land in New
South Wales as a recompense for losses he and his father had suffered in government

PROC. LINN. SOC. N.S.W., 111 (2), 1989

56 JOHN VAUGHAN THOMPSON, E.L:S.

service (Thompson, 1835)*. The Secretary of State replied that the regulations did not
allow him to make the grant (Colonial Office, 1835b).

To minimize his expenses, Thompson secured the post of Surgeon-Superintendent
on the emigrant ship Boadicea taking 200 unmarried females and about 60 people in
family groups to Van Diemens Land. For this service he received £50 and his own free
passage, but he had to pay what seems an exorbitant price of £159 for cabin passages for
his wife and children. On 8 August 1835 he signed his contract with the Secretary of the
Emigration Committee, undertaking to be available from 26 September ‘to inspect the
emigrants and witness their promissory notes’ (Colonial Office, 1835c). The Boadicea
sailed on 1 October 1835, and arrived in Hobart Town on 4 February 1836. It is an
extraordinary coincidence that Charles Darwin, who would later praise Thompson’s
work on barnacles, sailed into the Derwent River next day on the Beagle. There is no
evidence that Thompson and Darwin met, although they were in Hobart Town at the
same time for more than a fortnight.

There were no deaths on the 128-day voyage. The Surgeon-Superintendent
reported that: ‘I am quite confident that it was to [the] regulation of their diet, the with-
drawal of their wine while in the Torrid Zone and the promptitude with which every case
of sickness was met that under Divine Providence I am indebted for the preservation of
very many lives — not that we escaped our share of sickness nor the occurrence of
several very hopeless cases & almost miraculous escapes} (Colonial Office, 1836b).

After almost a month waiting in Hobart Town, Thompson and his family secured a
passage to Sydney on the North Briton, arriving on 15 March 1836 (Herald, 1836). He took
up his new duties in the Office of Deputy Inspector General of the Convict [Civil] and
Military Hospitals in the Colony of New South Wales on 1 April.

Sir James McGrigor, head of the Army Medical Department since 1815, was well
aware of the qualities of his senior officers, and proved a good friend to Thompson in all
that was to follow. He had sent Thompson his instructions on 20 August 1835 (Colonial
Office, 1835d). The Deputy Inspector General’s mission was to manage Convict and
Military Hospitals as a single service, as far as possible under the Hospital Regulations
of the Army. He was to give particular attention to keeping down costs, to controlling the
requisition and issuing of medicines and making regular returns about their use, and to
regulating the access by civil servants and other free residents to hospital services.

With uncanny prescience, McGrigor (Colonial Office, 1835d) advised Thompson
against possible pitfalls: ‘In the allotment of the duties civil and military you will take
care as much as possible to preserve the harmony necessary to good order; and in
assimilating the duties you will not unnecessarily disturb the Colonial Surgeons’
appointments, and on all occasions obtain the approbation of the Governor and proper
authorities in recommending any change of duty among inferior officers and servants’.

Early Administrative Problems

In ‘preserving harmony’ and ‘not disturbing the Colonial Surgeons, Thompson
failed utterly. Accordingly, his previous biographers have either discreetly ignored the
New South Wales part of his career or written about that to the exclusion of all else.

Part of the reason for his failure was undoubtedly the forthright and abrasive per-
sonality that he had already revealed in his scientific disputes and in his rejection of an
offensive complaint about rations from the sea-lawyers on the Boadicea (Colonial Office,

* This memorial has been mentioned already as the source of hitherto unknown information about
Thompson’s family and his service in Mauritius.

{ Im fact, it is the frankness and directness of Thompson’s reports on the emigrants that first attracted the
writer’s attention to him. He seemed so determined to face opposition head on, and to do it in writing,
(Campbell, 1988) that I was compelled to find out what happened to him afterwards — and before.

PROG. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL D//

1836d). In 1838, Governor Gipps said of him: ‘Dr Thompson, whatever may be his other
merits, wag not the man calculated to carry successfully into effect a measure In itself un-
palatable to his subordinates, being on the one hand wanting in blandness of manner or
conciliatory address, before which opposition might have gradually given way, and on
the other in that firmness and decision of purpose which would have overruled it’
(Colonial Office, 1838a).

But as MeGrigor wrote: “His situation was no easy one for any man of common
patience and temper. It was perhaps to be expected that an officer experienced in the
duties, sent out to control expenditure, diminish emoluments and supervise the manner
in which the several officers discharged their duties would be unacceptable, let his
address and manner be what they would’ (Colonial Office, 1839).

First, he had to serve three masters: the Governor and Secretary of State for the
Colonies in matters relating to the Convict [Civil] establishment, the Secretary at War
in matters relating to the Army, and the Director General of the Army Medical Service
in regard to professional reports and returns. What is more, running through his deal-
ings with all three was an undercurrent of resentment from officials in the Colonial
Office about the intrusion of the Army Medical Service and the War Office into their
domain. Hence, the Governor and his London superiors seemed to take some pleasure
in pointing out Thompson's failings to McGrigor, who was responsible for appointing
him and for disciplining him thereafter.

Second, through an incredible bureaucratic bungle, he arrived in Sydney to find
that Dr James Bowman, who had joined the colonial medical service in 1817 and had
been its head since 1819, was still in that position, and no arrangements had been made
to remove him from it. Consequently, one of the first recommendations Thompson had
to make was that his predecessor should be stood down. In the event, it took two years for
the despatches necessary to terminate Bowman’s appointment to pass back and forth
between Sydney and London. Meanwhile, Bowman remained on strength without
duties, devoting himself to his extensive private practice and the pastoral schemes of his
father-in-law, John Macarthur. Even without those supplements, his unearned salary
was some £200 greater than the pay for Thompson’s active Army rank.

Third, the other colonial surgeons, especially James Mitchell, who had been on
good terms with Bowman, also resented ‘Thompson’s arrival. His lack of a medical
degree did not go unnoticed, and the colonial surgeons were aware of his proposals to
replace them by army surgeons at lower rates of pay when their posts became vacant.
The security of their lucrative private practices was also threatened by their new lability
to transfer. And Thompson was less than tactful towards them. He was critical, in
reports to Sir James McGrigor, of their private practices, their drinking, and their
failure to cooperate with him.

Fourth, to cap Thompson's difficulties, the Governor reported at the end of 1836
that ‘Mr Thompson has not as yet presented any distinct and comprehensive arrange-
ment for. . . hospitals in this Colony, chiefly | apprehend from the infirm state of health
under which he has for some time been suffering (Colonial Office, 1836c).

In fact, Thompson had devoted a good deal of time to pointing out that his travel-
ling allowance did not cover the cost of the journeys he was required to make; comparing
his own pay and allowances with those of other senior officials; confronting the colonial
surgeons; and complaining to London about the obstacles the latter were putting in his
path.

‘To put the matter bluntly, the duties Thompson was expected to carry out, and the
circumstances in which he found himself, were beyond the powers of an elderly, ailing,
quarrelsome and forthright army surgeon who had been five years retired, and who in

PROG. LINN. SOG. N.S.W., 111 (2), 1989

58 JOHN VAUGHAN THOMPSON, F.L.S.

any case would rather have been working at his microscope. The consequences were
inevitable.

Nevertheless, Sir James McGrigor in London argued in his support: ‘Mr
Thompson complains that he has met with vexatious obstruction from some of the
Colonial Medical Officers at the outset; perhaps it 1s but natural that some of these
gentlemen should, at first, feel a new control irksome, and complain of more duty being
imposed on them than under the old system so as to interfere with lucrative private
practice or their avocations as agriculturalists . . .

‘I am not prepared in every instance to support the tone of Mr Thompson's
querulous correspondence, and I have so expressed my disapprobation of it that I trust
this zealous and experienced officer . . . will henceforth be most respectful in all his cor-
respondence . . . and I am of the opinion that henceforth no infirmity of temper will be
betrayed by the Deputy Inspector General of Hospitals.

‘In justice, however, to Mr Thompson I ought to say that during a long period of
service I have ever found him an able and honourable officer, one who has constantly
evinced the most indefatigable zeal on every service on which he has been employed,
acting with discretion, prudence and forbearance’ (Colonial Office, 1837a).

Despite this testimonial and promise for the future, in September 1837 a dispute
known as ‘the Mitchell case’ made public a conflict which had been festering since
Thompson’s arrival and which dragged on for almost four years. Briefly, Mitchell, who
had been in rancorous disagreement with Thompson from the outset, claimed to have
misunderstood an instruction, and so failed to attend as medical officer at a flogging.
Consequently the Governor dismissed him for disobeying an order. In response to pub-
lic support for Mitchell, Thompson inflamed the debate by writing a letter to a
newspaper; in it he referred selectively to some comments made about Mitchell by the
Governor. It is true that the Governor had more than once criticized the way the colonial
surgeons, including Mitchell, responded to Thompson’s instructions; but he had criti-
cized the tone and nature of the instructions themselves almost as often. Mitchell sued
Thompson for libel, and won. In the meantime, he had appealed against his dismissal.

The appeal led eventually to an inquiry by Governor Gipps: as a result, Mitchell
was reinstated in 1841 for one day, for the sole purpose of allowing him to retire without
a stain on his name. Although the finding supported Thompson’s rank and office, he
suffered more than Mitchell, who in fact had little to lose. But Thompson was left a
broken man, after yet another acrimonious conflict which had simply deepened his
bitterness and disillusionment.

An account of the quarrel between Thompson and Mitchell, distinctly unsym-
pathetic to Thompson, is provided by McIntosh (1956). It contains a good deal of accu-
rate detail but does not, in the writer’s opinion, sufficiently recognize the problems
Thompson faced in dealing with an entrenched Sydney establishment. A good
summary of the evidence is given in Gipps’ Report on his Review of the case (Colonial
Office, 1841b).

Thompson made other administrative mistakes, too. Soon after his arrival, he was
taken to task for writing personal letters to the Secretary of State for the Colonies about
mismanagement by certain officers of the colony who were outside his jurisdiction. He
was told to stop his ‘political investigations and writings’ (Colonial Office, 1837b), and
concentrate on his public duties. And as time went on, he displayed an increasing reluc-
tance to provide the reports and accountings required of him, leading to comments
about his ‘inflexible taciturnity’ and ‘habitual delay’ (Colonial Office, 1842a). What is
more, he did not bring about the hoped-for reduction in costs.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

HUGH CAMPBELL 59

Sustained Interest in Natural History

Despite Thompson’s difficulties with his official duties in New South Wales, he
continued to show his interest in natural history. Except for the dedication of his Zoologi-
cal Researches to Sir James McGrigor, he had scrupulously kept this interest separate from
his life as an army surgeon.

He began optimistically. On 20 April 1836, he wrote to his old acquaintance,
Alexander Macleay, the Colonial Secretary, expressing his disappointment at not find-
ing ‘the Society”™* flourishing, and his hope that Macleay would re-establish it and name
some original members. He also made a number of suggestions about the scope and
purpose of a natural history society, recommending a wide membership (Thompson,
1836). It appears that this letter was Thompson’s acceptance of an invitation to join ‘A
Committee of Superintendence of the Australian Museum and Botanical Garden’
(Gazette, 1836). Macleay became chairman, and Thompson was a member. The minute
books of the Committee show that he attended meetings regularly from 1836 to 1843
and was a member of the Sub-Committees for both the Botanical Gardens and the
Australian Museum (Australian Museum, 1836.)

Meanwhile, he still found time to look to his scientific reputation and contacts. His
letter to Julien Desjardins, President of the Natural History Society, Mauritius, claim-
ing authorship of the Catalogue of Exotic Plants in Mauritius has been mentioned already. A
letter from the Curator of the Australian Museum records that ten specimens of native
birds were given to J. V. Thompson ‘for the Mauritius’ in 1837 (Australian Museum,
1837). He is also on record in 1839 as having sent cotton seeds to Mauritius in 1838; and
the 1840 Annual Report of the Mauritius Natural History Society records that “Thomp-
son is the only one of our New South Wales correspondents who has not given us up’
(Vaughan, 1953). Thompson would have been heartened, too, to find that in 1839 Du
Cane, a friend of Macleay, was one of those who had confirmed his observations on the
metamorphosis of the Crustacea (Fletcher, 1920).

Removal from Office

Ever since Thompson had left Cork in 1835 without settling his account with Mrs
Dwyer, she had been trying to get her money. Failing in appeals directly to Thompson,
she had then unsuccessfully sought to have Sir James McGrigor order Thompson to pay.
Getting no satisfaction in that quarter either, on 28 October 1841 she complained to the
Secretary of State for the Colonies. It was Thompson’s bad luck that her letter arrived
only a fortnight after the Secretary of State had been considering Governor Gipps’s
report on the Mitchell case, and so Thompson's name was fresh in his mind. There is no
record to show that Thompson paid the bill, in spite of a fresh instruction to do so.

During 1842, Thompson’s neglect in submitting reports and returns became
intolerable to his superiors (Colonial Office, 1842b). Finally, in November, he was given
six weeks to submit a particular return. When he did not meet the deadline, preliminary
steps were taken to replace him (Colonial Office, 1843a). The Governor reported to the
Secretary of State, who in his turn asked McGrigor to remove Thompson. By this time it
was July 1843. In the letter to McGrigor, the Colonial Office reminded him of Mrs
Dwyer’s case, saying that ‘this renewed proof of that officer’s negligence’ seemed to call
for his suspension from duty (Colonial Office, 1843b).

McGrigor replied that he was already aware of the situation, and recognized the
need to act: ‘.. . for some time back I have had occasion to notice to Mr Thompson
many instances of neglect in conducting the public duties and of his rarely performing
the duty of inspecting the hospital, but finding this proceeded from greatly impaired

* Presumably he was referring to the defunct Philosophical Society of Australasia.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

60 JOHN VAUGHAN THOMPSON, FL.S.

health, with mental as well as bodily infirmity, I informed him of my intention. . . of his
being placed on the half pay of his rank in the army’ (Colonial Office, 1843c).

McGrigor followed up his official letter with a private one to Stephen, his counter-
part in the Colonial Office, presumably to make sure that no one proceeded with the
idea of suspending Thompson or dismissing him outright, and so cutting off his income:
‘The neglect of his duties by Mr Thompson .. . is not unknown to me. I regret to say
that for a considerable time back I have had cause to complain of the imperfect manner
in which he has performed his duties. I have reason to believe that this has proceeded
from the broken down health of Mr Thompson who is no longer equal to the multi-
farious duties of the responsible situation in which he is placed’ (Colonial Office, 1843d).

It is curious that Governor Gipps made no mention of Thompson's deteriorating
health, which had been mentioned by Bourke at the end of 1836 but ignored thereafter
in despatches from New South Wales. Hence one wonders how McGrigor came to his
conclusion that ill health was an important reason for Thompson’s poor performance.
But the question is academic: Gipps had secured Thompson’s removal.

Thompson was allowed to remain in office until his successor arrived in New South
Wales (Colonial Office, 1843e). He relinquished his position on 1 February 1844, and his
retirement to half pay was gazetted on 11 February (Johnston, 1917).

It should be said in his defence that, despite the problems described above, he sur-
vived for almost eight years in a position which had been almost untenable from the out-
set — more than twice as long as his counterpart in Van Diemens Land, who had taken
up duty at the same time and was superseded in 1839. And no one seems to have com-
plained about the quality of the medical service under his regime. In fact, at the end of
1839 Gipps had made a special point of remarking that he had no complaint about the
way the ‘strictly professional duties of the Medical Department in this Colony are
performed’ (Colonial Office, 1838b).

Thompson's ‘taciturnity’ and failing health in the 1840s did not prevent him from
returning to his old interest in botany, and four more papers from him on the cultivation
of cotton and sugar cane were published in the /ndian Journal of Agriculture between 1842
and 1845 (Royal Society, 1871).

When Thompson retired, one piece of business with the Colonial Office was still
unfinished. On 6 April 1843, having heard of a plan for sending young prisoners from
Parkhurst to the colonies for rehabilitation, Thompson wrote another characteristically
naive proposal to the Secretary of State for the Colonies. He announced that he had ‘just
purchased a farm (part of Eagle Farm) about seven miles from Brisbane ‘Town, and
[had] at present about 200 acres of clear tillage land’. His scheme was hastily conceived
and impracticable. He said he would be ‘most happy to try the experiment with from 10
to 20 of the Parkhurst boys . . . [T]hey will be removed from all temptation to steal or
pick pockets and will be employed in gardening, assisting the agricultural labourers and
vine dressers and in looking after the farm work. My object is to give a fair trial to tropi-
cal agriculture and in particular to cotton, sugar cane and coffee in addition to the vine
and other colonial productions’ (Thompson, 1843).

Perhaps Thompson wanted only to tell someone that he had just bought a property
to replace the land his father had lost sixty-five years earlier. Whatever he intended, his
plan misfired. It was typical of his luck that his application for Parkhurst boys was con-
sidered and rejected just the day before the Secretary of State approved his transfer to
half pay. His Lordship remembered the letter, and saw it as just one more proof of
Thompson’s inattention to his real duties (Colonial Office, 1843f).

Thompson’s last official letter is a sad reminder that mail took several months to
pass between London and Sydney. A week after Thompson had been retired to half pay,
the Governor called for a copy of his letter about the Parkhurst boys. Jno V. Thompson,

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 61

Deputy Inspector General, replied on 21 February 1844: ‘I have the honor to acknowl-
edge receipt of your communication of the 17th instant (44/14), and regret that I am
incapable of supplying the copy of the letter addressed to the Secretary of State of 6th
April last, considering it to be a private affair and only a request, which, having been
deprived of every means of accepting now by the loss of my appointment, I must of
necessity relinquish (Thompson, 1844).

As far as is known, Thompson remained in Sydney for the next three years, and so
did not have a chance to occupy his land at Moreton Bay. He died at his residence in
Liverpool Street on 21 January 1847, and was buried in the Parish of St Lawrence,
County of Cumberland (Registrar General, 1847). Presumably his grave was in the old
Devonshire Street cemetery, now the site of Central Railway Station.

He directed that the monies from his estate should be used in the first instance to
pay his debts, and that what remained should be paid to his ‘dear wife for her use
entirely (Supreme Court, 1847).

CONCLUSION

Since his death, John Vaughan Thompson’s contribution to natural science has
been recognized spasmodically. A crustacean, Vaunthompsonia, was named after him, as
was a plant from Madagascar, Dezdamia thompsoniana, though the latter was subsequently
given another name. He merits a line in some histories of zoology (Singer, 1959);
Lankester (1890) gives him a full paragraph in the Zoology article in The Advancement of
Science, remarking that “Thompson made ... great discoveries, which seem to have
fallen in his way in the most simple and natural manner, but must be regarded really as
the outcome of extraordinary genius’; and he 1s the subject of entries in the Dictionary of
National Biography and the Dictionary of Scientific Biography under his own name. His Zoo-
logical Researches were reissued in a facsimile edition in 1968. It is interesting that the
burning issues of his day were not seen by later biographers to be controversial. For
instance, metamorphosis in Crustacea, the subject of acrimonious debate in the 1830s,
was overlooked by Lankester, and hardly mentioned by Stebbing (1910) in the paper he
wrote to mark the centenary of Thompson’s election to a Fellowship in the Linnean
Society.

Today, however, 140 years after his death, the way he approached natural history is
more important than the truths he revealed. He was one of the genuine discoverers in
the age when gentleman naturalists played a significant part in scientific inquiry, and
even in that he illustrates a point: he shows how hard it was for a man short of money,
and working alone, to have his work noticed. Previous biographers have praised his per-
ceptiveness, and shown how his discoveries resulted from acute practical observation.
But they have not made enough of his persistence, his solitariness and the disappoint-
ments he suffered.

He was a ‘difficult’? man and yet he was able to make a notable contribution to both
botany and zoology. And that is only one side of his life. It is too easy to forget that his
eight-year stint in New South Wales, where the odds were against him from the first, was
the only blemish on a career in the Army Medical Service spanning forty-five years. On
balance, his unhappy inability to get on with people neither cancels out his work as a
surgeon nor dulls the brilliance of his lonely scientific achievements.

It is a pity that he was not in England when his discoveries were finally recognized.
There he might have been honoured instead of being ignored. The careful obituary in
the Sydney Herald of 26 January 1847, with no mention of friends and colleagues who
appreciated him, emphasizes his loneliness. And it was John Vaughan Thompson’s
usual luck that the age was wrong. He was in his 68th year, not his 63rd:‘. . . in his 63rd

PROG. LINN. SOG. N.S.W., 111 (2), 1989

62 JOHN VAUGHAN THOMPSON, EL.S.

year, after long-continued illness — distinguished for his achievements in zoology and
botany — possessing talents of no common order — and estimable in every relative duty
of life — he is deeply lamented by his afflicted family, to whom his loss is irreparable’
(Herald, 1847).

ACKNOWLEDGEMENTS

The Public Records Office, London, has granted permission for reproduction of
the Crown Copyright Colonial Office documents quoted in this paper. Acknowledge-
ment is made to the Australian Joint Copying Project of the National Library of Aus-
tralia and the Library of New South Wales for the microfilm of some of those
documents, and to the Archives Office of Tasmania and the State Library of Tasmania
for the remainder of the documents and for the facilities for reading all of them; to the
Archives Office of New South Wales, the Library of the Australian Museum, the
Library of the University of Edinburgh and the Library of the Linnean Society of
London for the references ascribed to them below; and to the Archives Office of the
Government of Mauritius.

Thanks are due to Guy Rouillard, President of the Historical Society of Mauritius,
Gina Douglas, Librarian and Archivist of the Linnean Society of London, P. H.
Sooprayen, Chief Archivist, Government of Mauritius, Jo Currie, Library Assistant,
Special Collections, University of Edinburgh Library, and Mary Kumvaj, Deputy
Librarian, Australian Museum. Each took the trouble to send me more information
than I asked for.

References

Key to abbreviations used in the text: ADB = Australian Dictionary of Biography, 1967; DNB = Dictionary of
National Biography, 1900; EB = Encyclopedia Britannica, 1968; Linn. Soc. = Linnean Society of London;
Zool. Soc. = Zoological Society of London.

Key to abbreviations used in the references: CO xx/xx = Colonial Office Item Numbers; AJCP xx =
Numbers of Australian Joint Copying Project Microfilm Reels; HRA = Historical Records of Aus-
tralia volumes. Otherwise standard abbreviations are used for societies and journals.

Since many of the references are to manuscript letters and other records, the references are presented in two
parts: Part 1: Published Works and Part 2: Unpublished letters, etc.

Part 1: References to Published Works

ADB, 1967. — Australian Dictionary of Biography. D. PIKE (ed.). Melbourne: Melbourne University Press.

BARNWELL, P. J., 1941. — Thompson, John Vaughan: Dictionary of Mauritian Biography, No. 3: 94-95.

BRITTEN, J., 1912. — John Vaughan Thompson. /. Bot. Lond. 50: 169-171.

CAMPBELL, H., 1988. — A proper class of female emigrants. Zasmanian Historical Research Association Papers and
Proceedings. Vol. 35, No. 2. A detailed account of the voyage of the Emigrant Ship Boadicea, J. V.
Thompson Surgeon-Superintendent.

DARWIN, C. R., 1851 — A Monograph on the Sub-class Cirripedia with figures of all the species. Lepadidae: 8. London:
Roy Society.

DNB, 1900a. — Vaughan, John: Dictzonary of National Biography, 58: 169. London: Oxford University Press.

——., 1900b. — Thompson, John Vaughan: Dictionary of National Biography, 56: 218. London: Oxford Univer-

sity Press.
EB, 1968. — Monmouth Court House, Battle of: Encyclopedia Britannica. Chicago: Encyclopedia Britannica
Inc.

ENCYCLOPEDIA AMERICANA, 1983. — Havana: Encyclopedia Americana. Danbury, Conn.: Grolier Inc.

FLETCHER, J. J., 1920. — The Society’s heritage from the Macleays. Proc. Linn. Soc. N.S.W. 45: 626-627.

GAZETTE DE LISLE Maurice, 1816. — Projected departures from Port Louis. Samedi 8 juin 1816. Mauritius.
Tome 2, page 2, cols. 3 & 4.

GAZETTE, 1836. — New South Wales Government Gazette. No. 226, 14 June 1836: 451.

HERALD, 1836. — Shipping Arrivals. Herald, Monday 21 March 1836, page 2. Sydney.

——,, 1847 — Obituary. Herald, 26 January 1847, page 3. Sydney.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

HUGH CAMPBELL 63

INTERNATIONAL GENEALOGICAL INDEX, 1981. — Index for Northumberland, C0664. International Genea-
logical Index. Salt Lake City: Church of the Latter Day Saints.

JOHNSTON, WILLIAM, 1917. — Roll of Commissioned Officers in the Medical Service of the British Army, 1727-1896,
H. A. L. HOWELL, (ed.): 132. Aberdeen: University Press.

LANKESTER, E. Ray, 1890. — The history and scope of Zoology. The Advancement of Science, Occasional Essays
and Addresses: 335-336. London: Macmillan.

MANNICK, A. R., 1979. — Mauritius, The Development of a Plural Society: 39. Nottingham: Spokesman.

McIntosh, A. M., 1956. — The case of Dr James Mitchell. Med. J. Aust. (5) August 4, 1956: 161-168.

ROYAL SOCIETY OF LONDON, 1871. — Thompson, John Vaughan. Catalogue of Scientific Papers 1860-1863, 5:
958-959. (Metuchen, N.J.: Scarecrow Reprint Corporation, 1968.)

SINGER, C., 1959. — A Short History of Scventific Ideas to 1900: 473. Oxford: Oxford University Press.

SLABBER, __, 1778. — Natural Amusements and Microscopical Observations. Haarlem. (Unidentified, apart from
mention in J. V. Thompson’s Memoir 1, 1828.)

STEBBING, T. R. R., 1910. — On John Vaughan Thompson and his Polyzoa.. . Proc. Linn. Soc. Lond., 123rd
session, 1910-11: 64-72.

THOMPSON, J. V., 1807a. — Catalogue of Plants growing in the vicinity of Berwick-on-Iweed. London. 1807. octavo,
132 pp.

—, 1808. — An Account of some new species of Piper [Pepper], with a few cursory observations on the
genus. [2 June 1807]. Trans Linn. Soc. Lond. 9: 200-203.

—., 1813. — Description of a new species of the genus Mus, belonging to the section of Pouched Rats. [1812.]
Trans Linn. Soc. Lond. 11, part 1 (20 April 1813): 161-163.

——, 1816 (claimed). — A Catalogue of Exotic Plants cultivated in the Mauritius. Mauritius: Baron & Souvignee.

——., 1827. — Memoir on Pentacrinus Europaeus, a recent Species discovered in the Cove of Cork. Cork. 4to, 2 plates.

——., 1828-34. — Zoological Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known
Animals. Cork. (A privately printed collection of six memoirs in five numbers issued between 1828 and
1834. Facsimile published London: Society for the Bibliography of Natural History. 1968. Introduc-
tion by Alwyne Wheeler.)

—_, 1828. — Memoir J: On the Metamorphoses of the Crustacea, and on Zoea, exposing their singular
structure and demonstrating that they are not, as has heen supposed, a peculiar Genus, but the Larva
of Crustacea!! Memoir 2: On the Genus Mysis, or Opossum Shrimp. Zoological Researches and Illustrations
or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 1. Cork.

—, 1829. — Contributions towards the natural history of the Dodo (Didus ineptus, Linn.) a bird which
appears to have become extinct towards the end of the seventeenth or the beginning of the eighteenth
century. Mag. Nat. Hist. 2: 442-448.

——, 1830a. — Memoir 4: On the Cirripedes or Barnacles . . . and the Class of Animals to which they belong.
Zoological Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 3.

Cork.

——,, 1830b. — Memoir 5: On Polyzoa, a new animal discovered as an inhabitant of some Zoophites . . . Zoo-
logical Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 4.
Cork.

VAUGHAN, R. E., 1953. — A forgotten work by John Vaughan Thompson Proc. Roy. Soc. Arts G Sez. Mauritius
1, part 3: 241-248.

vicors, N. A., 1830. — (In a review of Rathke’s just-published work on the development of the Crayfish).
Zool. J. 5, part 18 (June 30, 1830): 241-255.

WESTWOOD, J. O., 1835. — Of the supposed existence of Metamorphoses in the Crustacea. Phil. Trans Roy.
Soc. Lond. 125: 311-326.

WHEELER, A., 1970. — Thompson, John Vaughan: Dictronary of Scientific Biography 13. New York: Scribner.

——,, 1975. — Thompson: Marine Biologist. Brit. Med. J. 3: 534.

ZOOL. Soc., 1830. — Minutes, 28 December 1830. Proc. Zool. Soc. 1830: 140.

——,, 1831. — Minutes, 11 Jan. 1831 Proc. Zool. Soc. 1831: 144.

Part 2: References to Unpublished Letters and Miscellaneous Records

AUSTRALIAN MUSEUM, 1836. — Minute Books of ‘A Committee of Superintendence of the Australian
Museum and Botanical Garden’, 1836-1843. Australian Museum Archives. Sydney.

——., 1837. — Records of G. Bennett, Curator, 20 February, 1837. Australian Museum Archives. Sydney.

COLONIAL OFFICE, 1834. — CO 202/31 (AJCP Reel 222). Hay to Bourke, 18 December 1834.

——,, 1835a. — CO 201/251 (AJCP Reel 200). War Office to Stewart, Treasury, 15 June 1835.

——., 1835b. — CO 202/31 (AJCP Reel 222). Colonial Office to Thompson, 11 September 1835.

—, 1835c. — Archives Office of Tasmania, CSO 1/848/17942. Contract between Marshall and Thompson,
8 August 1835.

——,, 1835d. — CO 201/249 (AJCP Reel 200). McGrigor to Thompson, 20 August 1835.

——,, 1836a. — Archives Office of Tasmania, CSO 1/848/17942. Boadicea Passenger lists.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

64 JOHN VAUGHAN THOMPSON, F.L:S.

——., 1836b. — Archives Office of Tasmania, CSO 1/848/17942. Thompson’s Report, 4 February 1836.

——, 1836c. — CO 201/255 (AJCP Reel 203). Bourke to Glenelg, 10 November 1836.

——, 1836d. — Archives Office of Tasmania, CSO 1/848/17492. 14 passengers on the Boadicea to J. V.
Thompson, 20 November 1835.

——,, 1837a. — CO 201/269 (AJCP Reel 209). McGrigor to Stephen, 27 June 1837.

——., 1837b. — CO 201/268 (AJCP Reel 212). Thompson to Sec. of State, 23 September 1837.

——., 1838a. — HRA I, Vol. 19; 391. Gipps to Glenelg, 20 April 1838.

——., 1838b. — HRAI, Vol. 19: 396. Gipps to Glenelg, 24 April 1838.

——., 1839. — HRA, Vol. 20: 77. McGrigor to Stephen, 5 February 1839.

——, 1841a. — CO 201/313 (AJCP Reel 336). Mrs Dwyer to Sec. of State for Colonies, 28 October 1841.

——.,, 1841b. — CO 201/306 (AJCP Reel 331). Gipps to Sec. of State, 19 January 1841.

——.,, 1842a. — CO 201/321 (AJCP Reel 340). Gipps to Stanley, 20 June 1842, margin note.

——, 1842b. — CO 201/321 (AJCP Reel 340). Gipps to Stanley, 20 June 1842.

——, 1843a. — CO 201/331 (AJCP Reel 346). Colonial Sec. to Thompson, 7 January 1843.

——.,, 1843b. — CO 201/330 (AJCP Reel 346). Colonial Office to McGrigor, 22 July 1843.

——, 1843c. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 26 July 1843.

——, 1843d. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 1 August 1843.

——, 1843e. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 1 September 1843.

——,, 1843f. — CO 201/331 (AJCP Reel 346). Col. Office to Gipps, 7 September 1843.

EDINB. UNIv., 1797-8. — General and Matriculation Albums, 1797-8 and 1798-9. Special Collections,
University of Edinburgh Library.

——, 1816. — Institutions of Medicine Class Lists, 1816- 7. Special Collections, University of Edinburgh
Library.

LINNEAN SOCIETY of LONDON, 1807. — Thompson to Seaforth, 7 April 1807. The Linnean Society of
London Archives.

——.,, 1809. — Certificate of Recommendation for Fellowship of J. V. Thompson, signed 21 November 1809
by James Sowerby; Alex. Macleay, John Macleay and A. B. Lambert. The Linnean Society of London
Archives.

——.,, 1835. — Thompson to Linn. Soc. Treasurer, 8 May 1835. The Linnean Society of London Archives.

REGISTRAR GENERAL, N.S.W., 1847. — Deaths, 1847, Vol. 32. Registrar General of New South Wales.
Sydney.

SUPREME Court, 1847. — Probate, 19 February 1847. Supreme Court of New South Wales. Sydney.

THOMPSON, J. V., 1807b. — On the genus Kaempferca. Read to Linn. Soc., 7 April 1807.

——., 1835. — CO 201/251 (AJCP Reel 201). Thompson to Glenelg, Secretary of State for the Colonies,
7 August 1835.

—, 1836. — Archives Office of New South Wales, 4.2539.3, file 41/10460. Thompson to Macleay, 20 April
1836.

——, 1843. — CO 201/343 (AJCP Reel 354). Thompson to Sec. of State for Cols., 6 April 1843.

——., 1844. — HRAT, Vol. 23: 439, J. V. Thompson to Colonial Secretary Thomson, 21 February 1844.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

A List of the Cryptogams and Gymnospermous
Plant Specimens in the British Museum
(Natural History) gathered by Robert Brown
in Australia 1801-5

E. W. GROVES and D. T. MOORE

(Communicated by T. G. VALLANCE)

Groves, E. W., & Moore, D. T. A list of the cryptogams and gymnospermous plant
specimens in the British Museum (Natural History) gathered by Robert Brown in
Australia 1801-5. Proc. Linn. Soc. N.S.W. 111 (2), 1989: 65-102.

A list is here published for the first time of the cryptogams and gymnosperms
gathered by Robert Brown in Australia and now preserved in the herbarium of the
British Museum (Natural History), London. Brief remarks on Brown’s Australian visit
and collecting during 1801-5 are given. Correlation with the register of the collection
prepared by J. J. Bennett is indicated.

E. W. Groves, 143 Westlergh Avenue, Coulsdon, Surrey CR3 3AK United Kingdom (formerly
Department of Botany, British Museum (Natural History), and D. T: Moore, Department of
Mineralogy, British Museum (Natural History), Cromwetl Road, London SW7 SBD, United
Kingdom, manuscript received 12 April 1988, accepted for publication 15 February 1989.

. That full account of the Botanical discoveries
made during Flinders’s expedition, which the pub-
lic had a right to expect, has never appeared...’

J. Lindley, 1844

INTRODUCTION

The work of Robert Brown during and after Flinders’ expedition to Australia in
1801-5 is recognized as a significant contribution to Australian natural history in general
and Australian botany in particular, but scientific records of the expedition are frag-
mentary. Flinders’ (1814) account of the voyage of the Jnvestigator concentrates on geo-
graphical, navigational and sailing aspects and is useful for the chronology of the
expedition up to his departure in 1803 and subsequent imprisonment at Mauritius. But
for botanical details we must rely on Brown. His Prodromus of 1810 was intended to be the
main account of the botany, but only one volume of the two proposed was produced, and
only a few copies were sold. In due course, this important work was withdrawn. Brown
also contributed a modest botanical appendix to the Flinders volumes (1814: 2, 533-613).

After the death of the expedition’s commander, Matthew Flinders [1774-1814],
British interest in the voyage appears to have declined, and it was not until 1904 that
Britten published an account of the plants collected by Brown in Madeira on the out-
ward voyage. More recently Rourke (1974) has produced an account of Brown’s activi-
ties at the Cape of Good Hope, but there appears to be no list of South African
specimens collected by Brown in existence. Brown’s Timor plants are also little known,
being only briefly mentioned by Forbes (1885). In Australia Brown prepared lists of
plants seen or collected, or both, at some of the anchorages. Where these lists are still
extant they remain in manuscript, and as Lindley (1844) pointed out, no published list
of Brown's Australian plant specimens was available forty years after Brown returned. It

PROG. LINN. SOG. N.S.W., 111 (2), 1989

66 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

was not until the 1870s that a manuscript list of Brown’s specimens was prepared. This is
the catalogue referred to below as the ‘Bennett Register’.

Recent Australian interest in the voyage began with the account by Burbidge (1956)
of Brown’s nomenclatural system (based on Flinders’ chart annotations) for the Aus-
tralian collecting localities. A landmark in studies of Brown’s Australian botanical work
was reached a few years later when a facsimile edition of the Prodromus (Brown, 1810) was
published with an introduction by Stearn (1960a). There has been renewed interest in
the natural history aspects of the voyage in the United Kingdom recently with the publi-
cation of a paper on the expedition by Edwards (1976) and the publication of the diary of
Peter Good [d. 1803] also by Edwards (1981). Some of the paintings by the botanical
artist Ferdinand Lukas Bauer [1760-1826] are reproduced by Stearn (1960b), Stearn and
Blunt (1976) and Norst (in press). Vallance and Moore (1982) have given an account of
the geological aspects of the voyage and Mabberley’s (1985) work on Brown has provided
a much-needed biography. Clements (1983) has described the orchids collected by
Brown which are in the Lindley Herbarium and is to give an account of those in the
British Museum (Natural History) Herbarium. However, there are no published lists of
all the 3400 plants collected by Brown in Australia.

Further work, which will be published in the near future, includes the edited trans-
cription of Brown’s Diary of the voyage (Vallance et a/., in preparation). An important
outcome of the Diary publication project is the realization that considerably more speci-
mens were collected by Brown than are mentioned in either the Prodromus (Brown, 1810)
or the Diary accounts. In his Diary Brown uses a variety of plant names. These include
names already published by, for example, Carl Linnaeus [1707-1778], William Aiton
[1731-1793] and others. But he also used ‘nicknames’ for species he did not immediately
recognize. These are sometimes manuscript names attributable to D. C. Solander [1736-
1786]. Their use is hinted at in a letter (British Library Add. MS 32439, ff61-4 and
Fiistorical Records of N.S.W. 4: 776-9) written by Brown to Sir Joseph Banks [1743-1820]
from Sydney on 30 May 1802 which contains the following passage: ‘. . . The list. . . is
partly compos'd of nicknames given at the moment the plants were collected, and not
allow’d since. Some of them, I am afraid, may mislead, and the greater part I am aware
are rather barbarous, especially where I have terminated specific names in oides, which
I have done when uncertain whether the plant was distinct from a species already
known,...’

‘Oides’ names are mentioned in Brown’s Diary text and as a result of work
connected with editing the Diary for publication, the surviving plant specimens
gathered by Brown have been located in the BM (NH) Herbarium in order to check if
these ‘nicknames’ were mentioned on the original labels. Fortunately most of them are.
The locating of Brown’s plants led to the preparation of an index of over 3000 cards
(Groves and Moore, 1986), and the cryptogams and gymnospermous representatives
located in this way form the basis of this account.

ROBERT BROWN’S PLANT COLLECTION

A letter from Sir Joseph Banks to Sir John Barrow [1764-1848], written on 9th
October 1805 (B.L. Add. MS 32439, f185), reveals that twelve boxes of dried plants and
eleven boxes of Bauer’s natural history drawings were landed at Liverpool from the
Investigator. Vhere were, in all, 38 cases of natural history specimens (cf. Edwards, 1976).
Banks indicates in a letter of 3rd January 1807 (B.L. Add. MS 32439, ff237-245) that
about 3600 specimens were in the boxes of plants. Edwards (1976), citing Banks, stated
that there were about 700 plant specimens from the south coast of Australia, 500 from
the east coast, 500 from the north and 700 from Tasmania; there were some 1000

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 67

specimens from the neighbourhood of Port Jackson, and 200 from the island of Timor
(not discussed here).

Brown wrote relatively little on the lower plants, but the existence of his paper on
mosses (Brown, 1811) indicates he was to some extent interested in Bryophytes. But his
greatest contribution to the plants that are our concern in this paper was the realization
that the cycads had coniferous similarities (Brown, 1826).

Having survived the voyage to England in a leaking ship, impounding at the
Custom House of Liverpool, and a journey to London by the roads of 1805 (Vallance et
al., in prep.), 1t is appropriate to examine what befell the specimens after their arrival in
London. They appear to have gone initially to Banks’s house in Soho Square. There and
later at the British Museum, they were a major part of Brown’s herbarium (Murray,
1904). During his lifetime Brown kept his own herbarium separate from the British
Museum material and did not allow free access (Edwards, 1976), but George Bentham
[1800-1884] consulted it in the preparation of his Flora (Bentham and Mueller, 1863-
1878) (cf. Stearn, 1981). The next to feature as custodian of Brown’s plants was J. J.
Bennett [1801-1876]. Bennett became assistant to Brown at the British Museum in
November 1827 and Brown’s herbarium was left to him at Brown’s death (Woodward,
1904; Mabberley, 1985). In his turn Bennett kept the Brown herbarium separate from
the main herbarium but on his death (February 1876) his wife gave it to the British
Museum and it was then incorporated into the General Herbarium. Duplicates, and
material which today would not necessarily be considered as duplicate, were later dis-
tributed to other institutions in Europe and Australia. Those sent to Ireland have been
listed by Powell and Morley (1976).

Unlike the practice in other British Museum departments, the Botany Department
never prepared formal registers, so there are no present-day BM (NH) herbarium
numbers as such.

There are indications that some Brown Herbarium vascular plant specimens were
given to Banks’s librarian Jonas Dryander [1748-1810] during his lifetime. The Brown
specimens listed in the Bennett register do not include these duplicates. In our list we
have included them only where we believe the original is lost and where we can relate
them to the Bennett register entries. These Dryander duplicates are recognizable by
Dryander’s later label, but bear neither a Brown original label nor the blue sticker of
Bennett’s plants (see below), nor a number. Some are unique while others are plants
from the same gathering as that to which a Bennett number was given. The actual
numbering was done by James Britten [1846-1924], assistant and later librarian, in the
Botany Department and was probably undertaken soon after Brown’s herbarium was
acquired on the death of Bennett.

Not all species given by Brown (1810) are in our list, as Brown sometimes selected
Banks and Solander specimens for his descriptions. As an example, Mitrasacme prolifera
R.Br., Spigeliaceae (Bennett 2894) bears a Brown label indicating that it came from the
Endeavour River, Queensland, in 1770. The description of this taxon in the Prodromus
(Brown, 1810: 453, entry 8) then, is of a Banks and Solander gathering, not Brown's.

Fortunately, Brown annotated his algal and pteridophyte original labels after the
publication of Turner’s (1807-19) Fuci and his own Prodromus (Brown, 1810). So we have
reason for thinking that certain specimens are illustrated, or at least noticed, in these
works. A fern example is Bennett 104 (Gleichenia rupestris R.Br., see below). Bryophyte
and gymnosperm labels are not annotated in this way.

‘THE BENNETT REGISTER

After coming into possession of Brown’s Herbarium Bennett instituted a register of

PROC. LINN. SOC. N.S.W., 111 (2), 1989

68 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

the material. This gave the plants both a number and a name. But it was left to Britten,
assisted by Henry Trimen [1843-1896], to complete the task (Edwards 1976). No locality
or ecological data are recorded in this ‘Bennett Register. Such detail as Brown com-
mitted to paper is obtainable only from the original labels on the herbarium sheets, and,
in some instances, from Brown’s descriptions in the Botany Library (the ‘Brown Slips’).
In the herbarium the Brown specimens catalogued by Bennett and his followers are
recognized by a blue label bearing Britten’s hand-written number and the printed words
‘Iter Australiense 1801-5”.

One hundred and twenty-three numbers were allocated to Brown’s algae and
charophytes in the Bennett register. Of these, we have been able to find eighty-four.
Because Bennett’s workers missed specimens which we include, it is difficult to make
numbers of ‘found’ taxa agree with the Bennett register. Some of Brown’s algae were
described by Dawson Turner [1775-1858], and in the list below, dates of publication of
parts of Turner's Fuci (1807-1819) are given according to Price (1984). Fortunately, the
specimens described by Turner (1807-19) are recognizable because Brown annotated his
labels to the effect ‘this 1s the specimen sent to Mr Turner’.

The absence of Brown’s original labels in some cases means there is no locality
information for some specimens. Also, the Brown locality ‘South Coast’ is disappoint-
ingly vague. But better locality data are often available where a specimen is mentioned
in Turner (1807-19). In our list nomenclature, where appropriate, is in accordance with
Womersley (1967, 1984, 1987). The charophyte nomenclature used in our list is that of
Wood (1965) and we have located four out of five of Brown’s charophytes.

The lichens were never given space in the Bennett register. It would appear that
they were handed in the 1870s to the Rev. J. M. Crombie [1830-1906] to investigate,
probably with a batch of blank ‘Iter Australiense’ labels. It is Crombie’s writing on the
blue labels accompanying Brown’s original lichen labels and it would appear that
Crombie also applied the series of numbers. The numbers used are between 500 and
579 (thus not duplicating any numbers assigned by Bennett and Britten). It is these
numbers which were quoted by Crombie (1879). It can be seen from the lichen list that
there are several species given the same number. This is because since the 1870s many
have received revision and so now consist of more than one species under one number.
Many of Brown’s lichens were collected in Tasmania, and Kantvilas (1983) warns of the
unreliable nature of many of Brown’s labels. But, as others have found (Vallance and
Moore, 1982), the original labels may give reliable provenance data and should not be
regarded lightly.

Assuming that originally there were seventy-nine lichens, and each was allocated a
number by Crombie, we have been unable to trace twenty-four of them.

The fungi too are somewhat neglected by the Bennett register. They were allocated
numbers 142-156 (inclusive), but no identifications or other data were entered against
the numbers. The specimens, probably all mounted, were transferred to Kew Her-
barium when the BM(NH) fungal collection was passed to Kew as a result of the
Moreton agreement of 1961 (cf. Stearn, 1981: 309-310). They have since been incor-
porated into the Kew collections. A search made there by one of us (EWG) to trace
Brown’s fungi, with the aid of the list of the ten names in Brown’s appendix to Flinders
(1814: 2, 592-4), has revealed only one possible fungus. This is ‘Clavaria coralloides L’
(= Clavaria cristata var. coralloides Corner). While not attributed to Brown on the sheet,
the writing beneath the specimen and the watermark of the paper appear to be con-
temporary with other Brown sheets at the BM(NH).

The majority of Brown’s bryophytes in the BM(NH) remain unmounted and have
still to be critically examined. As they lack Bennett numbers they have not been in-
cluded here. But a minority were allocated Bennett numbers and subsequently

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 69

incorporated in the herbarium. These appear to be mostly from Tasmania. There is
Brown material on some sheets from the Hooker herbarium (ex Kew) but this does not
have original labels. From Brown’s Diary (cf. Vallance et al., in prep.) we know that
Brown visited only Port Dalrymple, and the Esk Rivers and the River Derwent/Mt.
Wellington areas of Tasmania in 1804. Despite this, we have accounted for all the spaces
in the Bennett register assigned to bryophytes.

In the 1870s Bennett assigned one hundred and forty-four numbers to Brown’s
pteridophytes. We have found all but two.

Brown (1810) indicated by letters the provenance of the species described, for
example ‘D’ for Tasmania, and ‘]’ for Port Jackson. The original labels were annotated
by Brown so there is reason for thinking that certain specimens are the ones that formed
the basis of the descriptions in Brown (1810).

Three cycadophytes are noted in the Bennett register. While we have traced two,
there remains uncertainty over the identity of ‘Zamia spiralis’ of Flinders (1814: 1, 81). It
appears to be a late entry to the Bennett register and thus existed in the 1870s. We have
not traced it. Brown (1810: 348, entry 1 ‘J’ & ‘M’) indicated his specimen was not tropical
and from study of Brown’s Diary (Vallance et a/., in prep.; cf. Flinders, 1814: 1, 348) we
believe the specimen came from Lucky Bay, Western Australia. Consequently, we think
it was, in fact, Macrozamia riedle: (Fisch. ex Gaudich.) C. A. Gardn., Zamiaceae (cf.
Johnson, 1959: 103). This 1s ‘Macrozamia spiralis Miq’ of Mabberley (1985: 86).

In our list of Brown’s conifers we follow the nomenclatural recommendation of Mr
K. Hill of the Royal Botanic Gardens, Sydney. Bennett assigned nineteen spaces in his
register to this group. We have found fourteen specimens.

The manuscript Bennett register (now housed in the Botany Library, BM (NH) ),
is the only document to list the plant specimens gathered by Brown in Australia. But it
was written seventy years after the voyage ended and is a somewhat inadequate docu-
ment for curatorial purposes. There were, for example, 3900 numbers allocated but con-
siderably fewer than this number of actual specimens; there are blank numbers in the
register and the numbers do not run consecutively. For example, the bryophytes finish at
295 and the next entry, the Asteraceae (Compositae), start at 2000. So, as the sequential
arrangement in the Bennett register bears no chronological relation to the order of
collection, and exhibits only a rudimentary grouping into families, there is no advan-
tage in giving our list in this manner. Consequently in our list the genera are arranged in
alphabetical order under their respective groups. The number we have given to each
entry has been applied merely to aid reference.

NOTES ON THE SPECIMEN LIST

1. In the following list we give Brown’s original label entry, with his geographical and
ecological data as they are set out on the labels. Keys to the geographical localities and
anchorages around Australia will be found in Burbidge (1956) and Stearn (1960a).

2. For some listed specimens there is more than one original label per herbarium
sheet, but in numerous instances there is more than one sheet per label and more than
one plant per sheet. In some cases it is not always clear which label goes with which
plant.

3. We have kept editorial comment on the texts of Brown’s original labels (given here in
single quotation marks) to a minimum, but we have indicated any references on the
original labels to Turner’s Fucz (1807-1819), Crombie (1879) and Brown (1810). The note
‘Icones Bauer’ found on some higher-plant labels appears not to be present on the
pteridophyte labels. Our claim that the Bauer painting of, for example, Azolla, is based
on a particular herbarium specimen is circumstantial and visual.

PROC. LINN. SOC. N.S.W., 111 (2), 1989

70 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

ALGAE

CHLOROPHYTA

1. Anadyomene browni (Gray) J.Ag. Bennett 205
Lacks a Brown original label, and is thus unlocalized. An annotation on the sheet
indicates that this is ‘Calomena brown1 of Gray (1866: t.44, fig. 3) and has type status.

2. Caulerpa browni (C.Ag.) Endl. Bennett 173
‘Actaia selago? [sic] Kents Islands’
There are two sheets of this material with identical labels.

3. Caulerpa cactordes (Turn.) C.Ag. Bennett 166
‘Actaia opuntia South Coast the specimen sent to Mr Turner’.
‘Fucus cactoides’ of Turner (1809-11: 3, 89-90, t.171). Type status.

4. Caulerpa flexilis Lam. Bennett 172
(Syn. ‘Caulerpa hypnoides’ (R.Br.) C.Ag.)

‘Actaia hypnoides Kents Islands’. Turner (1809-11: 3, 93-4, t.173).

Two specimens, one with type status.

5. Caulerpa geminata Harv. Bennett 167
(Syn. ‘Caulerpa sedoides’ (R.Br.) C.Ag.)

Two identical labels on the sheet. ‘Actaia sedoides L Kents Islands. The specimens sent
to Mr Turner’ In fact, ‘Fucus sedoides’ of Turner (1809-11: 3, 92, t.172). There is an
annotation on another sheet ex Turner herb. (formerly at Kew) by I. R. Price, 16 x11 1974
to the effect that the specimen was the basis of Turner’s figure. Type status.

6. Caulerpa racemosa var. laetevirens (Mont.) Weber van Bosse Bennett 168
No original labels, and specimen is consequently unlocalized, but a note by I. R. Price
of 10 x11 1974 indicates he considered the material to be possibly of the tropical var.
lamourouxit (Turn.) Weber van Bosse.

7. Caulerpa racemosa (Forsk.) J.Ag. var. laetevirens (Mont.) Weber van Bosse Bennett 169
‘Actaia clavata South Coast’. I. R. Price in a note of 10 x11 1974 considers this locality
incorrect.

8. Caulerpa racemosa (Forsk.) J.Ag. var. laetevirens (Mont.) forma cylindrica (Sonder)
Weber van Bosse Unnumbered
(Syn. ‘Caulerpa cylindracea’ Sonder)

Two specimens on one sheet, the lower one is labelled ‘Actaia clavata South Coast’

9. Caulerpa scalpelliformis (R.Br. ex Turn.) J.Ag. Bennett 171
Two labels: ist reads ‘Actaia scalpelliformis South Coast’; 2nd reads ‘Actaia
scalpeliformis South Coast the spec sent to Mr Turner’ Turner (1809-11: 3, 95-6, t.174).
Type status.

10. Caulerpa sertulariodes (S.Gmel.) Howe Bennett 170
(Syn. ‘Caulerpa taxifolia (Turn.) C.Ag.)

Two sheets, label reads ‘Caulerpa taxifolia Fucus taxifolius Turn South Coast’.

‘Fucus taxifolius’ is featured in Turner (1807-8: 1, 120-122, t.54). In a note on the sheet of
10 xi1 1974 I. R. Price considered this taxon to be tropical.

PROG. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 7

11. Caulerpa simpliciuscula (Turn.) C.Ag. Bennett 174
There are two sheets ‘Actaia conglobata Kents Islands. Turner 1809-11: 3, 98, t.175).

12. Codium australicum Silva Bennett 181 (pro parte)
No Brown label, consequently unlocalized.

13. Codium duthieae Silva Bennett 181 (pro parte)
No Brown label.

14. Codium fragile (Suringar) Hariot ssp. tasmanicum (J.Ag.) Silva Bennett 181
‘SC’ [?South Coast]. (pro parte)
15. Codium mueller Kutz. Bennett 181a

No Brown label and unlocalized.

16. Polyphysa peniculus (R.Br. ex Turn.) C.Ag. Bennett 203
(Syn. ‘Acetabularia peniculus’ (R.Br. ex Turn.) Solms-Laubach)
Has type status but sheet on loan in November 1988.

PHAEOPHYTA

17. Acrocarpia paniculata (Turn.) Aresch. Bennett 213
Two labels: Ist reads “The sp sent to Mr ‘Turner Fucus scoparino Kents Islands Bass

Strait’; 2nd reads ‘Fucus paniculatus ‘Turn. hist 3 p.99 t.176 Cystoseira paniculata
Agardh [1823-8] Sp. alg. 1 p.76. Turner (1809-11: 3, 99-100, t.176). Type status.

18. Asperococcus bulbosus Lam. Bennett 179
‘F[ucus] ampullaceus is an incorrect name for a plant wch when perfect is filld with a
dense gelatinous material’ [reverse reads] ‘Kents Islands & South Coast’.

19. Carpoglossum confluens (R.Br. ex Turner) Kutz. Bennett 226
‘Fucus confluens Port Dalrymple’. ‘Fucus confluens’ of Turner (1809-1811: 3, 17, t.141),
Agardh (1823-8: 1, 95). Type status.

20. Cystophora brown (Turn.) J.Ag. Bennett 208
‘Fucus Browni Turn Hist 4 pl t 197 Cystoseira Brownii Agardh sp algar; 1 p.73’ [reverse]
‘sent to Mr Turner, you proposd to give it my name South coast frequent. According to
Turner (1811-19: 4, 2-3), the specimen is from King George Sound. See also, Agardh
(1823-8: 1, 73). H. B. Womersley appended a note, dated 31 v 1952, to the effect that the
lower of the two specimens on the sheet is the type of ‘C. browni’, but the upper may be
‘C. monilifera’.

21. Cystophora moniliformis (Esper.) Womersley & Nizamuddin ex Womersley

Bennett 214
Two labels, 1st reads ‘non logica F barbata sed distincta & prope F thuyoid: Herb Banks’
[reverse] ‘Fucus paniculatus litt. Kents Group’; 2nd reads ‘Fucus spartioides Turn hist. 4
p.85 t.232. Mr Turner did not return the specimen which is in fructification RB’ Turner
(1811-19: 4, 86, t.232). Type status.

22. Cystophora retroflexa (Labill.) J.Ag. Bennett 209
Two labels: 1st reads ‘Fucus ramosissimus South Coast sent to Mr Turner’ 2nd reads

PROC. LINN. SOC. N.S.W., 111 (2), 1989

72 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

‘Fucus retroflexus Labill. Turn. hist. 3 p.47 t.155 Cistoseira retroflexa Agardh [1823-8]
sp Alg. 1 p. 7#. Turner (1809-11: 3, 48, t.155).

23. Cystophora torulosa (R.Br. ex Turn.) J.Ag. Bennett 211
‘Fucus torulosus Kents Islands Bass Strait 1803-4 The sp sent to Mr Turner. Turner
(1809-11: 3, 51, t.157). Another sheet (ex Kew herbarium), lacks a Bennett number, but is
almost certainly another portion of the same gathering. It has a label which reads ‘dup
Fucus torulosus N Sp Turn fuc 3 [. . .] t157 Kents Group Cystophora torulosa Agardh’.
Turner (1809-11: 3, 52, t.157). Type status.

24. Cystophora sp.indet. Bennett 219
‘Fucus vagus the sp sent to Mr Turner Fucus vagus South Coast. The Turner illustration
implied by the original label is untraced.

25. Cystophora sp.indet. Bennett 221
‘Cystoseira (?) affin retroflexa Dr Agardh has a perfect specimen from Dr Greville.
Unlocalized.

26. Cystophora sp.indet. Bennett 224
‘Cystoseira |. . .] retroflexa. Unlocalized.

27. Cystosezra trinodis (Forsk.) C.Ag. Bennett 218
(Syn. ‘Cystophyllum muricatum’ (Turn.) J.Ag.)

Two labels: Ist reads ‘Fucus angustifolius North Coast of New Holland F muricatus Mr
Turner’s writing’; 2nd reads ‘Cystoseira muricata North Coast Cystoseira trinodis Ag

[1823-8], Sp Alg p.67 [. . .] 24 June 1833’ Turner (1808-09: 2, 107-8, t.112).

28. Cystosezra trinodis (Forsk.) C.Ag. Bennett 223
No original labels, and unlocalized.

29. Dictyopterts woodwardia (R.Br. ex Turn.) J.Ag. Bennett 177
Two identical labels which read ‘Fucus woodwardia North Coast [reverse reads] a F
membranaceo divers frondibus spinulo-denticula. A third sheet ex Herb. Kew, although
lacking the Bennett number, is of the same gathering. Type status.

30. Dictyota sp.indet. Bennett 251
‘Kents Islands’.

31. Ecklonia biruncinata (Bory) Papenfuss Bennett 229
(Syn. ‘Ecklonia radiata (Turn.) J.Ag.)

‘Fucus radiatus ‘Turn. hist 2 p.161 t.134 Port Dalrymple too young for Baptism [sic].
Turner (1808-09: 2, 161-2, t.134).

An annotation on the sheet indicates this specimen was identified by H. B. S. Womersley
25. v. 1952 as the type of Turner’s var. exasperata of F. radiatus.

32. Homoeostrichus sinclairit (Hook. & Harv.) J.Ag. Bennett 175
(Syn. ‘Zonaria stuposa R.Br. ex J.Ag.)
Two labelled sheets: 1st reads ‘Zonaria Novy sp’; 2nd reads ‘Kents Islands’. Type status.

33. Hormosira banksit (Turn.) Decaisne No Bennett number
No sheet bearing Brown’s original label exists. There is, however, a label in Brown’s

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE the)

hand ex Herb. Kew which reads ‘Nov. Holl. or occident. Herb. Mus. Paris Sep 1816
[added later] Cystoseira expansa Ag’ [a third sheet is labelled] ‘Cistoseira Banksii. Type
status.

34. Macrocystus pyriforme (L.) C.Ag. Bennett 231
‘Fucus pyriformis King’s Island’.

35. Pertthalia caudata (Labill.) Womersley Bennett 227
Two labels: 1st reads ‘Fucus inermis Port Dalrymple & South Coast of New Holland;
2nd reads ‘Fucus inermis distinct from F aculeatus with minute globules terminating the
ramuli. Type status.

36. Phyllospora comosa (Labill.) C.Ag. Bennett 230
Two identical labels reading ‘Fucus polyphyllus South Coast of New Holland F
comosus. We note ‘Turner (1809-11: 3, 18, t.142) gives the locality as ‘Van Diemens Land’.

37. Platythalia quercifolia (R.Br. ex Turn.) Sond. Bennett 217
Three labels: 1st reads ‘Fucus quercifolius South Coast New Holland 1802 [. . .]; 2nd
reads ‘Fucus quercifolius South Coast of New Holland Turn hist 3 p37 t.151 Cystoseira
quercifolia Ag’; 3rd label reads ‘Fucus quercifolius Turn hist 3. p.37 t.151. Turner (1809-
11: 3, 37, t.151). Type status.

38. Sargassum decipiens (Turn.) J.Ag. Unnumbered
Two labels: Ist reads ‘Fucus polymorphus [struck through] decipiens Port Dalrymple the
sp sent to Mr Turner’; 2nd reads ‘Fucus decipiens Ag Turn fuc 3 p79 t166. Turner (1809-
bE 9> t166). Iivpe status:

39. Sargassum decurrens (Turn.) J.Ag. Bennett 157
‘Fucus decurrens Turn hist 3 pl43 t194 North Coast (I believ) [sic] only specimen sp
figured. See Turner (1809-11: 3, 143, t.194). Type status.

40. Sargassum heterophyllum C.Ag. Unnumbered
Turner (1808-9: 2, 62, t.92) cites a Brown specimen of ‘Fucus heterophyllus’ from ‘Coast
of New Holland’ but this material is untraced.

41. Sargassum lacerifolium (R.Br. Turn.) J.Ag. Bennett 158
‘Fucus lacerifolius Turn hist 3 p81 t.167 Sargassum lacerifolium Agardh [1823- 8] sp alg.
1- p15 [. . .]. From Port Dalrymple, Tasmania, according to Turner (1809-11: 3, 81-2,
t.167). Type status.

42. Sargassum paradoxum (R.Br.) Harv. Bennett 210
Two labels: Ist reads ‘Fucus paradoxus South Coast’; 2nd reads ‘Fucus paradoxus Mss
Turn Fuc 3 p.49 t156 Cystoseira paradoxa Agardh [1823-8] in sp. alg. 1 p.15. Turner
(1809-11: 3, 49, t.156). Type status.

43. Sargassum vestitum (R.Br. ex Turn.) C.Ag. Bennett 159
Two labels: 1st reads ‘Fucus vestitus Kents Islands Bass strait & South Coast The speci-
men sent to Mr Turner; 2nd reads ‘Fucus vestitus Turn. hist. 3 pl01 t177 Sargassum
vestitum Agardh [1823-8] Sp. Alg. 1 p24. Turner (1809-11: 3, 101-2, t.177). Type status.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

74 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

44, Sargassum sp.indet. Bennett 161
‘Sargassum North Coast of New Holland’.

45. Sargassum sp.indet. Bennett 162
‘Sargassum cum S. ilicifolia receptaculis convenit North Coast’

46. Sargassum sp.indet. Bennett 163
‘Sargassum North Coast of New Hollnad’

47. Scytothalia dorycarpa (Turn.) Grev. Bennett 216
Two sheets and two labels: 1st reads ‘Fucus platycarpus South Coast of New Holland
The specimen sent to Mr Turner; 2nd reads ‘Fucus dorycarpus Turn hist 3. p.21 t.143
Cystoseira dorycapa Agardh [1823-8] Sp alg 1 p.80°.

‘It is not clear in Turner (1809-11: 3, 24, t.144) ifthe illustrated specimen was collected by
Menzies or Brown.

48. Seirococcus axillaris (R.Br. ex Turn.) Grey. Bennett 215
Two sheets, three labels in all: 1st reads ‘Fucus axillaris Port Dalrymple The sp sent to
Mr Turner sp fig’; 2nd reads ‘Fucus axillaris Turn hist 3 p.27 tl46 Cystoseira axillaris
Agardh [1823-8] sp. alg p.80’; 3rd reads “Cystoseira axillaris Port Dalrymple’. ‘Turner
(1809-11: 3, 28, t.146). Type status.

49. Splachnidium rugosum (L.) Grev. Bennett 225
Two labels: 1st reads ‘Fucus rugosus Kents Islands shallow water on rocks. You say [sic]
Ulva rugosus Herb/?] Lad’; 2nd reads ‘Fucus rugosus Turn. hist 3 p.119 t.185 Agardh
[1823-8] Sp alg. 1 pl100 Kents Islands’ Turner (1809-11: 3, 118, t.185).

50. Sporochnus radiciformis (R.Br. ex Turn.) C.Ag. Bennett 228

‘Fucus ?radiciformis is ita Fucus Kents Islands Bass Strait. Type status.

51. Xiphophora chondrophylla (R.Br. ex Turn.) Montague ex Harv. Bennett 271
Two labels: ist reads ‘Fucus chondrophyllus Port Dalrymple’; 2nd reads ‘Fucus
chondrophyllus Mss Turn. hist 4 p. 60 t.222” Turner (1811-19: 4, 60, t.222).

52. Zonaria sp.indet. Bennett 182
No Brown label and consequently unlocalized.

RHODOPHYTA

53. Brongniartella australis (J.Ag.) Schmitz Bennett 195
‘Griffithsia australis Ag [. . .]. Unlocalized. Agardh (1823-8: 2, 135).

54. Champia parvula (C.Ag.) Harv. Bennett 240
No orig. label, consequently unlocalized.

55. Coelarthrum muellert (Sonder) Borges. Bennett 245
‘Fucus compar Kents Islands Bass Strait & South Coast of New Holland’. Another sheet
ex Herb. Kew, of the same gathering, bears the locality in Brown’s handwriting as ‘South
& East coasts of New Holland’.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 75

56. Gelidiella acerosa (Forsk.) J. Feldmann & Hamel Bennett 264
(Syn. ‘Gelidiopsis rigida’ Grev. )

‘Spharococcus rigidus Ag [1823-8] Sp. Alg. p.285 [. . .]. No locality or other data on
original label.

57. Gelidium australe J.Ag. Bennett 250
‘F corneus? [sic]. No locality given.

58. Gelidium glandulaefolium Hook. & Harv. Bennett 272
No Brown label. Secondary label reads ‘Fucus Kents Group’.

59. Gracilaria edulis (S.Gmel.) Silva Bennett 269
(Syn. ‘Gracilaria corniculata (R.Br. ex Turn.) J.Ag.)

Two labels: Ist reads ‘Fucus ceratophyllus [struck through] corniculatus The sp sent to
Mr Turner Kents Islands 1803-4 proxe cornea a distinc: spec? [sic]; 2nd reads ‘Fucus
corniculatus Turn 3 p.113 t.182. Turner (1809-11: 3, 112, t.182). A note appended by
H. B. S. Womersley on 27.v.1952 indicates ‘this specimen may well be a Gigartina. ‘Type
status.

60. Griffithsia sp. indet. Bennett 193
‘Griffithsia. No locality given.

61. Hymenocladia divaricata (R.Br.) Harv. Bennett 254
‘Fucus divaricatus Mss in fructific only specimen Fucus divaricatus Turn hist 3 p.110
t181. Turner (1809-11: 3, 110, t.181) localizes Brown’s material as from ‘on the coast of New
Holland’ ‘Type status.

62. Hymenocladia usnea (R.Br.) J.Ag. Bennett 259
‘Fucus usnea Mss Kents Group Basses Strait 1803-4. A second label, not Brown’s, reads
‘Fucus usnea Ag Turner Hist 4 p70 t225. Chondria usnea Agardh [1823-8] Syn Alg.
XVIIT. Turner (1811-19: 4, 70, t.225). Type status.

63. Hypnea musciformis (Wulf.) Lamour. Bennett 236
‘F musciformis. Unlocalized.

64. Laurencia botryowdes (Turn.) Gaill. Bennett 253
On loan at time of compilation.

65. Laurencia pinnatifida (Huds.) Lam. Bennett 257
Two sheets with identical labels; ‘Fucus pinnatifidus Kents Islands’.

66. Leptophyllis conferta (R.Br. ex Turn.) J.Ag. Bennett 263
Two labels: ist reads ‘Fucus confertus Kents Islands 1803 Scopula nov sp? [sic]’; 2nd
reads ‘Fucus confertus Mss Turn. 3 p177 t184 Delesseria ?conferta Ag. [1823-8] Sp. alg.
p. 177? [sic]. Turner (1809-11: 3, 116, t.184). Type status.

67. Melanthalia obtusata (Labill.) J.Ag. Bennett 260
Two labels: ist reads ‘Fucus strictus Port Dalrymple F obtusatus Labill’; 2nd reads
‘Fucus obtusatus Labill Turn. hist 3 p.25 t.145. Turner (1809-11: 3, 25-6, t.145).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

76 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

68. Osmundaria prolifera Lamour. Bennett 206
(Syn. ‘Polyphacum proliferuny C.Ag.)

‘Polyphacum proliferum Agardh [1823-8] Sp. alg. 1. p.106. Osmundaria prolifera
Lamour Port Dalrymple the only specimen’

69. FPeyssonnelia australis Sond. Bennett 176
No Brown label, consequently unlocalized.

70. Phacelocarpus labillardiert (Mert. ex Turn.) J.Ag. Unnumbered
No Brown gathering of this taxon has been traced in the BM (NH) herbarium, but there
is a specimen marked ‘Australia R Brown’ (not in Brown’s writing) and a fragment on
another sheet labelled ‘F scolopendrum Mss Mr Brown 1807 [sic]. Both sheets were
formerly in the Kew herbarium. ‘Turner states regarding ‘Fucus labillardierr that “This
plant has also been gathered upon the coast of New Holland by our own countryman,
Mr Brown’ Turner (1809-11: 3, 8, t.137).

71. Plocamium sp.indet. Bennett 251
‘Fuci pulchilli var? [y] Junior planta? Kents Islands Bass Strait according to you may be
F aspenoides in herb’ Secondary label reads “Delessaria plocamum [a].

72. Plocamium sp.indet. Bennett 252
‘Fucus pulchellus Kents Island in Bass Strait & Port Dalrymple 1803-4.

73. Polyopes constrictus (Turn.) J.Ag. Bennett 268
‘Two labels: Ist reads ‘Fucus densus Kents Islands 1803-4’; 2nd reads ‘Fucus constrictus
Ain: hist 3 p39 tla27 See lurmer (1809-115 3) 39=40) tlb2)) type status:

74. Folysiphonia decipiens Mont. Bennett 235
(Syn. ‘Polysiphonia cancellata Harv.)
Lacks original label, and consequently unlocalized.

75. Pterocladia lucida (R.Br. ex ‘lurn.) J.Ag. Bennett 266
‘Fucus lucidus Mss ‘Turn. hist. 4 p99 t238. ‘Turner (1811-19: 4, 99, t.238) gives the locality

as ‘on the southern coasts of New Holland’.

76. Rhabdonia verticillata Herv. Bennett 244
‘Fucus 1s it difft from Compar [sic] Kents Islands’.

77. Rhodymenia cunerfolia (Hook.f. & Harv.) Taylor Bennett 249
(Syn. ‘R. corallina’ Grey.)

No Brown label, consequently unlocalized.

78. Trematocarpus concinnus (R.Br.) De ‘Toni Bennett 233
‘Fucus caespitosus according to you a var. of F plicatus Kents Islands One spec in fructif.
‘Type status.

79. Trematocarpus concinnus (R.Br.) De Toni Bennett 267
Two labels: 1st reads ‘Fucus scoparius [struck through] concinnus Kents Islands 1803-4;
2nd reads ‘Fucus concinnus Mss ‘turn. hist 3 p41 t153. See Turner (1809-11: 3, 41-2,
t.153). Type status.

PROG. LINN. SOG. N.S.W,, 111 (2), 1989

E. W. GROVES AND D. T. MOORE 77

80. Vidalia fimbriata (R.Br.) J.Ag. Bennett 261
Two labels: Ist reads ‘Fucus fimbriatus North Coast of New Holl spec unic (?){sic]’; 2nd
reads ‘Fucus fimbriatus Mss ‘Turn. Hist. 3. p87 t170’. Turner (1809-11: 3, 87-8, t.170).
Type status.

CHAROPHYTA
81. Chara corallina Klein ex Willd. Bennett 277
(Syn. ‘Chara australis’ R.Br.)
‘Chara crassifolia Port Jackson’
82. Chara corallina Klein ex Willd. Bennett 279

(Syn. ‘Chara plebeja R.Br.)
‘Chara A Carpentaria Point [sic, locality not given].

83. Nitella congesta(R.Br.) A.Br. Bennett 276
‘Chara cfr B a Bay I South Coast’. Lucky Bay, Western Austalia (cf. Burbidge, 1956).

84. Nitella hyalina (DC.) J.Ag. Unnumbered
Two sheets with identical data ‘Chara B Point T Carpentaria. Cape Shield, Northern
Territory (cf. Burbidge, 1956).

LICHENES

85. Cladia aggregata (Sw.) Nyl. Bennett 532
‘Lichen multiflorius Cladonia multiflora ericet elevat aust:’ (cf. Crombie, 1879: 393), a
type according to Filson (1981: 15).

86. Cladia ferdinand (Mull. Arg) R. Filson Bennett 533
‘Lucky Bay Cladia retipora. Another sheet without a Bennett number, but of the same
gathering, bears the label ‘On the ground Lucky Bay Tasmania [sic]. A note by R.
Filson of 8 vi 1983 states that ‘this specimen was probably collected at Lucky Bay, W
Australia [...]’. The identification of this material has been confirmed by D. J.
Galloway as Cladia ferdinandit.

According to Crombie (1879: 393) Brown’s ‘Lichen cribrosus’ was from ‘Mountain
heaths, Australia’.

87. Cladta retipora (Labill.) Nyl. Bennett 534
‘Lichen cribrosus Cladonia cribrosa enceta Montes que ora australis & Ins Diemen’

Tasmania, cf. Crombie (1879: 393) and Filson (1981: 23).

88. Cladia retipora (Labill.) Nyl. Bennett 534
No original label, probably another part of the gathering above.

89. Cladonia squamosa Mill. Arg. Bennett 531
(Syn. ‘Cladonia subsquamosa’ (Nyl.) Wain var. ‘pulverulenta (R.Br.) Vain. and ‘C.
pulverulenta R.Br.)

No original label; Crombie’s handwriting reads ‘on the ground Table Mt. Tasmania R.
Brown Cladonia acuminata (Achy.

Brown’s ‘Cladonia pulverulenta was from Mt Wellington (cf. Crombie 1879: 392).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

78 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

90. Cladonia pleurota (Florke) Schaerer Bennett 530
(Syn. ‘Cladonia deformis’ Hoffm.)

‘lat. sax. Mont. Tabul prom Dieman’ Brown’s ‘Lichen deformis’ was from Mt Welling-
ton, Tasmania, cf. Crombie (1879: 393).

91. Cladina confusa (R. Sant.) Follm. & Ahti Bennett 527
‘Lichen rangiferinus mons Tabul & encet’.. Crombie (1879: 393) gives ‘Mt Wellington on
the ground.

92. Coenogonium implexum Nyl. Bennett 504
‘Lichen spongiosus arb cort: laevi in sylvis umbrosus humidis ad radis Mont Tabul
prom Diemen’. Mt Wellington, Tasmania (cf. Crombie, 1879: 399).

93. Collema durvetzu Degel. Bennett 511
‘Lichen fascicularis? rip & arb Musc insula Diemen’. This specimen had been earlier
identified by Crombie as ‘Collema implicatum’ Nyl. From trees near the River Derwent,
‘Tasmania according to Crombie (1879: 391).

94. Collema leucocarpum Hook.f. & Tayl. Bennett 549 (bis)
‘Lichen obductus rup & arb orce australis N Holl & Ind Dieman’. Crombie (1879: 391-2).
See also entry below.

95. Collema subconveniens Nyl. Bennett 549 (bis)
We have not been able to trace this specimen, but Degelius (1974: 139) appears to have
seen and cited it ‘as C. nigr[esens] Lichen obductus (BM) N.W. Tasm’ According to
Crombie (1879: 392) Brown’s ‘Lichen obductus’ was from trees near the River Derwent,
‘Tasmania.

96. Diuploschistes ocellatus (Vill.) Norman Bennett 563
‘Lichen lateritus [. . .] prope Risdon Cove’. Tasmania (cf. Crombie, 1879: 399).

97. Dirinaria picta (Sw.) Clem. & Shear Bennett 546
No original label; secondary label reads ‘Lichen pictus Sw near Kingstown, Newcastle’

(cf. Crombie, 1879: 397).

98. Ephebe fruticosa Henssen Bennett 522
Two species on sheet: label reads ‘Lichen pubescens Lin Ach L. exilis Lightf rup & sax
inundat’. Specimen from the Grose River, NSW, according to Crombie (1879: 391).

99. Ephebe tasmanica Cromb. Unnumbered
According to notes on the sheet there are three species present. The first is Ephebe tas-
manica Cromb. The second Ephebe fruticosa A. Henssen. The third is Pseudephebe pubescens
(L.) M. Choisy. (Syn. ‘Electoria pubescens(L.) Howe). According to Crombie (1879:
391) ‘Lichen pubescens’ was from River Grose, NSW.

100. Flavoparmelia rutidota (Hook.f. & Tayl.) Hale Bennett 539
‘Lichen cfr L caperatum fl Derwent. Tasmania (cf. Crombie, 1879: 394).

101. Graphis intricata Fée Bennett 561
‘Lichen Opegrapha dendritica arb: trunce rip: fl Grose [. . .]. According to Crombie

PROG. LINN. SOG. N.S.W,, 111 (2), 1989

E. W. GROVES AND D. T. MOORE 79

(1879: 401), the specimen was from the bark of trees, near the River Grose, New South
Wales.

102. Graphis platycarpa (Eschw.) A. Zahlbr. Bennett 560
‘Lichen Opegrapha plebja cor: arb: prope P Jackson. (Graphis sophistica Nyl. and
Opegrapha plebja of Crombie, 1879: 401).

103. Haematomma puniceum (Ach.) Mass. var. collatum (Stirt.) Zahlbr. Bennett 569
(Syn. ‘Haematomma collatum (Stirt.) Dodge)

‘Ad cortices Port Jackson. Probably “Lichen guttatus’ gathered from bark of trees near
Middle Harbour, Port Jackson; cf. Crombie (1879: 399).

104. Haematomma puniceum (Ach.) Mass. var. infuscum (Stirt. ex Bailey) Zahlbr.
(Syn. ‘Haematomma infusum (Stirton ex Bailey) R. W. Rogers) Bennett 507
‘Lichen guttatus Middle Harbour Port Jacksor (cf. Crombie, 1879: 399).

105. Hypogymnia lugubris (Pers.) Krog Bennett 526
‘Lichen nem aff L physodi lat Mont Tabularis’. Mt Wellington, Tasmania (cf. Crombie,
187925395):

106. Hypogymnia mundata (Nyl.) Rassad. Bennett 550 (bis)
According to an annotation by J. A. Elix (of x 1977) this specimen, noted by Crombie as
‘Parmelia mundatum var. pulverata Nyl. [. . .] Lichen dendrosme on trees base of Table
Mt, Tasmania, is a mixture of the above species and H. pulverata (Nyl. ex Crombie) Elix.

107. HAypogymnia pulverata (Nyl. ex Cromb.) Elix Bennett 515 (on some sheets as 550)
No original label, but has been written up by Crombie (1879) as a mixture of Parmelia
mundata Nyl. (ex Derwent River) and “Parmelia angustata’ Pers. (ex Port Jackson).
Remounting seems to have removed P mundata, and the latter has been identified by
J. A. Elix (x 1977) as the above.

108. HAypogymnia tabularis (Taylor) Elix Bennett 550
‘Lichen dendrosma [...] in sylvis umbrosis ad radic Montis Tabularis prope fl
Derwent. Determined by Crombie (1879: 395) as ‘Parmelia mundata Nyl., more
recently by J. A. Elix (ix 1977) as the above.

109. Lecidea flindersii Cromb. Bennett 565
‘Superce [?] terra Risdon Cove’. Tasmania (cf. Crombie, 1879: 400).

110. Melaspilea corcumserpens Nyl. Bennett 566
‘Lichen ambiguens I terra nude rarius in saxis In collibus prope Risdon Cove’ Tasmania

(cf. Crombie, 1879: 401).

111. Nephroma cellulosum (Ach.) Ach. Bennett 543
‘Lichen antarcticus rup umbrosos humid adrip fl Anna Maria infra calaratumnY

‘Anna Maria River appears to be Margate Rivulet, southeastern Tasmania, (cf.
Vallance et al., in prep.).

112. Pannaria rubiginosa (Thunb. ex Ach.) Del. Bennett 556
‘South coast Bay 3”. Fowlers Bay, South Australia (perhaps Crombie, 1879: 397).

PROC. LINN. SOG. N.S.W., 111 (2), 1989

80 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

113. Parmelia tenuirima Hook.f. & Tayl. Bennett 537
Two sheets: Ist reads ‘Lichen amplissimus in saxis & rupibus collium prope fl: Derwent’
2nd sheet has no orig. label. “Lichen amplissimus’ of Crombie (1879: 394).

114. Pseudephebe pubescens (L.) M. Choisy Bennett 521
‘Lichen lanatus sum Mont Tabul prom Diemen (cf. Crombie, 1879: 395).

115. Pseudocyphellaria billardiere: (Delise) Rasanen Bennett 536
‘Lichen linearis in trunc arbor prope fluv Derwent Tasmania (cf. Crombie, 1879: 396).

116. Pseudocyphellaria crocata (L.) Vainio Bennett 540 (bis)
‘Lichen crocatus Dicks: Hoffm: rup & Sax in montosis’ (cf. Crombie, 1879: 396). See P
neglecta, below, for second label.

117. Pseudocyphellaria glabra (Hook.f. & Tayl.) Dodge Bennett 538
‘Lichen latissimus rup & sax Montis Tabularis. Mount Wellington, Tasmania (cf.
Crombie, 1879: 396). Determined as ‘Sticta freycinetii’ Del. by Crombie.

118. Pseudocyphellaria multifida (Nyl.) D. Galloway & P. James Bennett 516
‘Lichen dichotomus rup: umbros: |. . .] ad rep fl Anna Maria prope Calaractam (cf.
Crombie, 1879: 396). Perhaps Margate Rivulet, Tasmania.

119. Pseudocyphellaria neglecta (Mull. Arg.) H. Magn. Bennett 540 (bis)
‘Lichen crocatus. Locality as 116 above?

120. Psoroma decipiens (Hedw.) Hoffm. Bennett 567
(2Syn. ‘Lecidea decipiens’)
‘Lichen decipiens [. . .] supra terra Risdon Cove’ Tasmania (cf. Crombie, 1879: 400).

121. Psoroma asperellum Ny)l. Bennett 547
‘Lichen adscendens [. . .] ad latus Mont: Montis Tabularis. Mt Wellington, Tasmania
(cf. Crombie, 1879: 398).

122. Psoroma hypnorum (Vahl) S. F. Gray Bennett 354
‘Lichen cfr: L hypnorum lat sax: Montis Tabularis prope Derwent’. Mount Wellington,
Tasmania.

123. Psoroma leprolomum (Nyl.) Rasanen Bennett 553
No original label, secondary label suggests specimen is ‘Lichen denrosmae’ from base of
Mt Wellington, Tasmania, cf. Crombie (1879: 398).

124. Psoroma soccatum R.Br. ex Crombie Bennett 502
‘Lichen soccatus trunce emiort ad radi Mont Tabularis’ (cf. Crombie, 1879: 398-9).
Lectotype (Galloway, 1985: 481).

125. Psoroma sphinctrinum (Mont.) Nyl. Bennett 553
‘Lichen Dendrosma reniles Derwent River’ Tasmania (?Crombie, 1879: 398).

126. Ramalina inflata (Hook.f. & Tayl.) Hook.f. & Tayl. Bennett 519
‘Lichen fastigiatus arb [. . .] prope P Jacksor’ (cf. Crombie, 1879: 393).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 81

127. Ramalodium succulentum Nyl. ex Cromb. Bennett 551
‘Lichen succulentus arb: trunc & rup umbros ad rip fl Grose [. . .] 2 rami and musci (cf.

Crombie, 1879: 392).

128. Relicina limbata (Laurer) Hale Bennett 544
Two specimens, but one original label missing; secondary label suggests ‘Lichen pin-

natus on rocks from Port Jackson. 2nd (original) labels reads ‘Lichen pinnatus in saxis
prope P Jackson’ (cf. Crombie, 1879: 394).

129. Siphula coriacea Nyl. Bennett 501
‘Lucky Bay S Ct. Another sheet, without a Bennett label, bears a label in Crombie’s
hand ‘on the ground Lucky Bay ‘Tasmania [sic].

See also number 86, above. Crombie (1879: 393), however, has number 501 from Mt
Wellington, Tasmania.

130. Sphaerophorus melanocarpus (Sw.) DC. Bennett 517
Two sheets: 1st label reads ‘Lichen fragilis arb trunc semi putrid ad rip flu alpinor Mon
Tabul prom Diemen’?; 2nd has no original label (cf. Crombie, 1879: 392).

131. Sphaerophorus tener Laurer Bennett 524
‘Lichen globiferus lat sax Mont Tabul prom Diemen (cf. Crombie, 1879: 392).

132. Stereocaulon ramulosum (Sw.) Rauschel Bennett 526
‘Lichen ramulosus men alt L paschali lat Mont Tabul &c rip fl Grose &c’ (cf. Crombie,
1879: 392).

133. Stricta subcaperata (Nyl.) Nyl. Bennett 555
‘Lichen filix Lin: fil. In lat umbros supra summitato Montis Tabularis insul. Dieman’.
Mount Wellington, ‘Tasmania (cf. Crombie, 1879: 396).

134. ‘Teloschistes chrysophthalmus (L.) Th. Fr. Bennett 545
‘Lichen chrysophtphalamus [. . .] Kents Group.’ Bass Strait (cf. Crombie (1879: 396-7).

135. Thamnolia vermicularis (Sw.) Ach. ex Schaerer Bennett 528
‘Lichen vermicularis Cladonia vermicularis lat & sum: Mon Tabul prom Diemen.
Mount Wellington, Tasmania (cf. Crombie, 1879: 393).

136. Thelotrema lepadinum (Ach.) Ach. Bennett 520
‘Lichen ? [sic] occulatatus Cort laevi: arbor ad ripas fluv: Grose [. . .]’ (cf. Crombie,

1879: 399).

137. Unmbilicaria cylindrica (L.) Delise in Duby Bennett 558
Two specimens: Ist has no original label, secondary label suggests ‘Lichen proboscideus
summit of Table Mtn? 2nd (original) reads ‘Lichen proboscideus [.. .] sum: Mont
Tabularis prope fl Derwent. Mount Wellington, Tasmania (cf. Crombie, 1879: 397).

138. Usnea acromelana Stirt. var. decipiens (Lamb) Lamb Bennett 523
(?Syn. ‘Neuropogon acromelanus’)
‘Lichen ustulatus rup sum: Mont Tabularis prom Diemen’ (cf. Crombie, 1879: 395).
Holotype of variety (Walker, 1985).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

82 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

139. Usnea ceratina Ach. Bennett 518 (bis)
(cf. Crombie, 1879: 394). Specimen untraced.

140. Usnea florida (L.) Wigg. Bennett 518 (bis)
‘Lichen floridus’ from the River Grose, N.S.W., specimen untraced. (cf. Crombie, 1879:
393).

141. Xanthoparmelia australiensis (Cromb.) Hale Bennett 525
‘Scyphophorus lat Mont Tabul In Diem [sic]’. Mt Wellington, Tasmania according to
Crombie (1879: 395), but this locality could be an error (Kantvilas 1983). According to
Bibby (1951), and Ehix et al. (1986) the locality is the Nullarbor region and Flinders
Ranges. Holotype of species (Elix et al., 1986).

142. Xanthoparmelia multipartita (Cromb.) Hale Bennett 542
‘Lichen multipartitus in saxis prope P Jackson’. (cf. Crombie, 1879: 394-5). There is
other material in the gathering with no original label; secondary label reads ‘Lichen
multipartitus rocks near Port Jackson’

Taxon distributed in south-eastern Australia and this number is described as type by
Elix et al. (1986).

BRYOPHYTA

LEAFY LIVERWORTS

143. Jamesoniella colorata (Lehm.) Spruce ex Schiffn. Bennett 293
Secondary label reads “Iasmania. ‘The taxon 1s known to occur in Tasmania according to
Scott (1985).

144. Frullania falciloba Yaylor ex Lehm. Bennett 290
Secondary label reads “Tasmania. Known in Tasmania (Scott, 1985).

145. Gackstroema weindorfi (Herzog) Grolle Bennett 294
Secondary label reads “Tasmania. Known to occur, frequently in Nothofagus forests, in
Tasmania (Scott, 1985).

146. Lepicolea scolopendra (Hook.) Dum. ex Trev. Bennett 292
Secondary label reads “Tasmania.

147. Plagiochila sp.indet. Bennett 291
Secondary label reads “Tasmania.

‘THALLOID LIVERWORTS

148. Hymenophyton flabellatum (Labill.) Dum. ex Trev. Bennett 295
(Syn. ‘Symphyogyna flabellata’ (Labill.) Mont.)

Secondary label reads “Iasmania. Known in ‘Iasmania, Scott (1985). The specimen is
unmounted, and in a folded and re-used sheet of Brown’s herbarium drying paper on
which is written ‘Carpentaria Island s[. . .] 21 Jany 1803 [. . . but this is struck through
in pencil.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 83

149. Symphyogyna podophylla (Thunb.) Mont. & Nees Bennett 296
(Syn. ‘Symphyogyna obovata Hook.f. & Tayl.)

Secondary label reads “Tasmania. The label is annotated ‘Syn Hep p.480’ (cf. Gottsche et
al., 1844-7).

Scott (1985) confirms the Tasmanian occurrence of this taxon.

MUSCI

150. Blindia magellanica Schimp. Bennett 284
Secondary label reads “Tasmania. Tasmanian occurrence is confirmed by Scott and

Stone (1976).

151. Breutelia affinis (Hook.) Mitt. Bennett 288
Secondary label reads “Tasmania (cf. Scott and Stone, 1976).

152. Breutelia pendula (Sm.) Mitt. Bennett 289
Secondary label reads “Tasmania (cf. Scott and Stone, 1976).

153. Bryum billardiert Schwaegr. Bennett 287
Secondary label reads “Tasmania (cf. Scott and Stone, 1976).

154. Campylopus clavatus (R.Br.) Wils. Bennett 282
Secondary label reads “Tasmania (cf. Scott and Stone, 1976).

155. Campylopus introflexus (Hedw.) Brid. Bennett 281
Secondary label reads “Tasmania (cf. Scott and Stone, 1976).

156. Dawsonia polytrichoides R.Br. Unnumbered
Ist original labels reads: ‘Nova Cambria Australis: ad radices montium prope Port
Jackson Jan 1 1804 RB [sic]’; 2nd original label reads ‘Dawsonia polytrichoides to show
that part of the peristomium originates from the columnale or Placenta. But an anno-
tation on the wrapping paper of an unmounted specimen in a ‘Herb. Brown’ cover reads
‘Genus Muscor frondos Proxim: Polytricho lesser division of the first branch of the
River Grose Jan 1 1805 desc’.

The 1st original label is incorrectly dated. Brown was in Tasmania 1 January 1804 (cf.
Vallance et al., in prep.). The date 1 Jan 1805 is consistent with the New South Wales
locality. Probably material from this gathering was described by Brown (1811) where the
locality was given as ‘Port Jackson. Scott and Stone (1976) note the occurrence of this
taxon in Tasmania, Victoria, New South Wales, A-C:T. and Queensland.

157. Dhicranoloma dicarpum (Nees) Par. Bennett 285
Secondary label reads “Tasmania (cf. Stone and Scott, 1976).

158. ‘Dicranoloma robustum’ of BM (NH) Herbarium Bennett 283
Secondary label reads “Iasmania’.

159. Leptostomum erectum R.Br. Unnumbered
Secondary label reads ‘New Holland Mr Brown’. The mounted Hooker specimen (ex
Kew Herbarium) bears a secondary label which reads ‘New Holland Mr Brown’. Taxon
known in Victoria, New South Wales and Queensland (Scott and Stone, 1976).

Possibly gathered with Brown’s (1811) material from the ‘Hawkesbury and Grose’.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

84 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

160. Leptostomum inclinans R.Br. Bennett 286
Possibly this material was gathered with that described by Brown (1811) and from ‘Van
Diemans Land’ A Hooker specimen of this gathering (ex Kew Herbarium) has a second-
ary label which reads “Table Mountain Van Diemens Land Mr Brown at an elevation of
3000-3500ft. A label on the sheet indicates this specimen 1s the basis of the illustration
by Brown (1811: 320, t.23, fig 2). A secondary label with the unmounted material reads
“Tasmania (cf. Scott and Stone, 1976).

161. Leptostomum macrocarpum (Hedw.) Pyl. Unnumbered
Secondary label reads ‘Leptostomum macrocarpum, Br Australia R Brown. Known
from Tasmania, New South Wales and Queensland according to Scott and Stone (1976).

162. ‘ayloria octoblepharum (Hook.) Mitt. Bennett 280
Secondary label reads “Tasmania. Scott and Stone (1976) confirm the occurrence in
Tasmania.

PTERIDOPHYTA

LYCOPSIDA

163. Lycopodium deuterodensum Herter Bennett 126 (on same sheet as Bennett 127)
‘Lycopodium candelabrum Derwent Huon &c (Lycopodium densum of Brown, 1810:
165, entry 3-D).

164. Lycopodium deuterodensum Herter Bennett 127 (on same sheet as Bennett 126)
‘Lycopodium venustrum Port Dalrymple Jan 1804 [reverse] Lycopodium candelabrum’.

165. Lycopodium deuterodensum Herter Bennett 128
‘Lycopodium candelabrum B & S Port Jackson [reverse] 3 Lycopodium densum’
(Brown, 1810: 165, entry 3-J).

166. Lycopodium fastigiatum R.Br. Bennett 129
Two labels, ist reads ‘4 Lycopodium fastigiatum Summit of Table Mountain VDL
(Brown, 1810: 165, entry 4-D); 2nd reads ‘Lycopodium quadrifarium Port Jackson’.

167. Lycopodium laterale R.Br. Bennett 130
‘Lycopodium diffusum Port del Esperance V D Land’ (Brown, 1810: 165, entry 5-D).

168. Lycopodium laterale R.Br. Bennett 131
‘Lyopodium laterale [. . .] Port Jackson’ (Brown, 1810: 165, entry 6-J).

169. Lycopodium myrtifolium Forst.f. Bennett 123
‘Lycopodium varium [alpha]|[. . .] Table Mountain’ (Brown, 1810: 165, entry 1-D).

170. Lycopodium myrtifolium Forst.f. Bennett 124
‘Lycopodium varium [beta] alpinae Mont Tabule’ (Brown, 1810: 165, entry 1-D). Holo-
type of ‘Lycopodium varium [beta] alpinum’, according to an annotation on the sheet by
J. M. Beitel 1981.

171. Lycopodium scariosum Forst.f. Bennett 125
‘Lycopodium decurrens affr L reptans B & S: Mont Tabularis and fluy Derwent’
(‘Lycopodium decurrens’ of Brown, 1810: 165, entry 2-D).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 85

172. Selaginella ulrginosa (Labill.) Spring Bennett 133

‘Lycopodium quadrilineum var Port Phillip’

173. Selaginella uliginosa (Labill.) Spring Bennett 132

‘Lycopodium uliginosum Labill’ (Port Jackson, cf. Brown, 1810: 165, entry 7-J).
PSILOPSIDA

174. Psilotum nudum (L.) Beauv. Bennett 120

Two labels: 1st reads ‘Lycopodium nudum Broad Sound [reverse] 1 Psilotum
triquetrum’; 2nd reads ‘Lycopodioides nuda P.Jacksor (‘Psilotum triquetrum’ of Brown,
1810: 164, entry 1-J).

175. Timesipteris billardiert Endl. Bennett 122 (on same sheet as Bennett 121)
‘Lycopodioides truncata [. . .]| Derwent [reverse] 2 Psilotum truncatum’ (Brown, 1810:
164, entry 2-D).

176. Tmesipteris truncata R.Br. Bennett 121 (on same sheet as Bennett 122)
‘Lycopodioides truncata [. . .] Port Jackson [reverse] 2 Psilotum truncatum’ (Brown,

1810: 164, entry 2-J).

FILICOPSIDA (TRUE FERNS)

177. Acrostichum aureum L. Bennett 2
Two labels: 1st reads ‘Acrostichum fraxinifolium prod 145 Arnhem South Bay Feb 6
1803’, Caledon Bay, Northern Territory (Burbidge, 1956); 2nd reads ‘Acrostichum fraxi-
nifolium [. . .] Port II prod 145’, Port Clinton, Queensland (Burbidge, 1956). Brown was
there in August 1802 (Vallance et a/., in prep.; Brown, 1810: 145, entry 2-T).

178. Adiantum aethiopicum L. Bennett 69
‘Adiantum assimile Shoal Water Bay Keppel Bay prodr 155.’. A. aethiopicum of Beadle et
al. (1972), perhaps Brown (1810: 155, entry 4). Brown, however, does not note a tropical
specimen (“I”).

179. Adiantum aethiopicum L. Bennett 70
‘Kangar|oo] Island’ (Brown, 1810: 155, entry 4-M).

180. Adzantum aethiopicum L. Bennett 71
‘Adiantum ebeneum Port Jackson’ (Brown, 1810: 155, entry 4-J).

181. Adiantum formosum R.Br. Bennett 67
‘Adiantum formosum prodr 155 Port Jackson’ (Brown, 1810: 155, entry 3-J).

182. Adiantum formosum R.Br. Bennett 68
‘Adiantum cfr formosum Port Jackson [. . .]’ (Brown, 1810: 155, entry 3-J).

183. Adiantum hispidulum Sw. Bennett 64
‘Adiantum hispidulum Shoal Water Bay prodr 155’ (Brown, 1810: 155, entry 2-T).

184. Adiantum hispidulum Sw. Bennett 65
‘Adiantum [. . .] hispidulosa Port Jackson’ (Brown, 1810: 155, entry 2-J).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

86 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

185. Adiantum hispidulum Sw. Bennett 66
Three labels: 1st reads ‘Adiantum cfr A. hispidulum Port Jackson; 2nd reads ‘North
rocks 8 August 1802’; (Brown, 1810: 155, entry 2-J). 3rd reads ‘Dove Dale Feb 1804 —’.

186. Arthropteris tenella (Forst.f.) J.Sm. Bennett 12
‘Polypodium tenellium Forst prodr 147 Polypodium fimiculus rupibus & rare in loc
umbrosis in rip[as] Grose’ (Brown, 1810: 147, entry 8-J).

187. Asplenium aethiopicum (Burm.f.) Bech. Bennett 33
(Syn. ‘Asplenium furcatum’ Thunb.)
‘Asplenium praemorsum prodr 150 rup umbrose: Grose’ (Brown, 1810: 150, entry 7-J).

188. Asplenium attenuatum R.Br. Unnumbered, but corresponds to Bennett register
entry 25

No original label; secondary label reads ‘Nova Cambria Australis: Port Jackson R

Brown (Brown, 1810: 150, entry 2-J).

A duplicate of this in Kew Herbarium, as No 26, is from the ‘Grose’ River, Blue

Mountains.

189. Asplenium bulbiferum Forst.f. Bennett 35
Two labels: Ist reads ‘Asplenium |. . .] laxum prodr 151’; 2nd reads ‘Asplenium laxum
prodr 151 (Brown, 1810: 151, entry 10-D).

190. Asplenium flabellifolium Cay. Unnumbered, but corresponds to Bennett register 30
‘Asplenium flabellifolium Van Diemens Land’ (Brown, 1810: 150, entry 6-D).

191. Asplenium flabellifolium Cay. Unnumbered but corresponds to Bennett register 31
‘Asplenium flabellifolium Port Jackson’ (Brown, 1810: 150, entry 6-J).

192. Asplenium nidus L. Unnumbered but corresponds to Bennett register 25 (bis)
‘Asplenium nidus Port Jackson’ (Brown, 1810: 150, entry 1-J).

193. Asplenium nidus L. Unnumbered but corresponds to Bennett number 25 (bis)
Three labels: Ist reads ‘Asplen. nidus North Coast’; 2nd reads ‘Asplenium nidus Linn
prodr 150 Broad Sound West Peaked Hill In sylvis umbrosis ad radices arborum spec
parasiticum” [cf. Brown, 1810: 150, entry 1-T]; 3rd reads ‘Grose 1804 (cf. Brown 1810:
150, entry 1-J).

194. Asplenium obtusatum Forst.f. Bennett 27
No original label; secondary label reads ‘Insula Van Diemen 1804 (probably Brown,
1810: 150, entry 3-D).

195. Asplenium paleaceum R.Br. Unnumbered but corresponds to Bennett register 29
No original label, secondary label reads ‘Nova Cambria australis in monte prope Broad
Sound 1802 Robert Brown’ (cf. Brown, 1810: 150, entry 5-T).

196. Asplenium polyodon Forst.f. Unnumbered but probably corresponds to
(Syn ‘A. falcatum’ Lam.) Bennett register 28
No original label; secondary label reads ‘Nova Cambria australis Port Jackson’ (perhaps
corresponds with Brown, 1810: 150, entry 4-J).

PROC. LINN. SOG. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 87

197. Azolla pinnata R.Br. Bennett 134
Two labels: 1st reads ‘Azolla pinnata Nob in stagnis prope Richmond in Noy Camb:
durt’; 2nd reads ‘Azolla pinnata Nob in stagnes & fluv len: fluat Richmond Hawkesbury’
(Brown, 1810: 167, entry 1-J).

Bauer painting 229 (BM (NH) Botany Library) is probably based on this plant or the
specimen below.

198. Azolla pinnata R.Br. Bennett 135
‘Azolla filiciloies [sic] pinnata Patersons River Oct 1804 (Brown, 1810: 167, entry 1-J).
BM (NH) Bauer painting 229 perhaps based on this plant, but see above.

199. Azolla filiculoides Lam. var. rubra (R.Br.) Strasburger Bennett 136
‘Azolla rubra Patersons River Oct 1804 (‘Azolla rubra’ of Brown, 1810: 167, entry 2-J).

200. Blechnum camfieldi Vindale Bennett 49
‘Blechnum mutatum var. Port Jackson 8 Stegania procera prodr 153’ (‘Stegania procera
of Brown, 1810: 153, entry 8-J).

201. Blechnum ambiguum (Presl.) Kaulf. ex C.Chr. Bennett 41
Two sheets, with identical labels ‘3 Blechnum laevigatum Prodr 152 Port Jackson’
(Brown, 1810: 152, entry 3-J).

202. Blechnum cartilagineum Sw. Bennett 39
‘Blechnum paludosum Port Jackson [. . .] 1 Blechnum cartilagineum (Sw) prodr 152’
(cf. Brown, 1810: 152, entry 1-J).

203. Blechnum chambersi Tindale Bennett 42
‘2 Stegania lanceolata prodr 152 Van Diemens Land’ (Brown, 1810: 152, entry 2-D).

204. Blechnum fluviatile (R.Br.) E. J. Lowe Bennett 43
‘Pteris fluviatilis rivul alp Mont tabul. Derwent [. . .] 3 Stegania fluviatilis prodr 152’ (cf.
Brown, 1810: 152, entry 3-D).

205. Blechnum indicum Burm.f. Bennett 40
‘2 Blechnum striatum prodr 152 Port Jackson’ (Brown, 1810: 152, entry 2-J).

206. Blechnum minus (R.Br.) Ettingsh. Unnumbered but probably corresponds
‘Insula Van Diemen 1804 RB’ (Brown, 1810: 153, entry 7-D). to Bennett register 47
207. Blechnum nudum (Labill.) Mett. ex Luerss. Bennett 45

‘Blechnum pteroides Port Dalrymple Jan 1804 5 Stegania nuda prodr 153’ (‘Stegania
nuda’ of Brown, 1810: 153, 5-D).

208. Blechnum nudum (Labill.) Mett. ex Luerss. Bennett 46
‘Blechnum ?discolour Osmunda discolor B&S Kings Id 6 Stegania falcata prodr 153
[. . .] (Stegania falcata’ of Brown, 1810: 153, entry 6-D).

209. Blechnum penna-marina (Poir.) Kuhn Bennett 44
‘Pteris alpina desc cat Mont Tabul Derwent 4 Stegana alpina prodr 152 (‘Stegania
alpina’ of Brown, 1810: 152, entry 4-D).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

88 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

210. Blechnum penna-marina ssp. alpina (R.Br.) Kuhn Unnumbered but

probably correponds to Bennett register 44
‘Insula Van Diemen in summitate Montis Tabularis 1804 RB Stegania alpina prodr
[. . .| (probably another sheet of the above, cf. Brown, 1810: 152, entry 4-D).

211. Blechnum watts Tindale Bennett 48
‘Blechnum mutatum desc Kings Island 8 Stegania procera prodr 153’ (Brown, 1810: 153,
entry 8-D).

212. Blechnum watts Tindale Bennett 49 (bis)
‘Port Jackson 8 Stegania procera prodr 153’ (cf. “Stegania procera of Brown, 1810: 153,
entry 8-J). The label bears an annotation by T. C. Chambers dated 6. x11. 1977 which
reads ‘Maybe a hybrid but closest to Blechnum watts Tindale.

213. Botrychium australe R.Br. Bennett 119
‘1 Botrychium australe prodr 164 Paramatta Port Jackson (Brown, 1810: 164, entry 1-J).

214. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 74
‘Adiantoides crispa Kangar[oo] Id’.

See Quirk et al. (1983: 510-3) regarding the relationship of Chezlanthes austrotenuifolia and
C. tenuifolia.

215. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 76
‘Adiantoides crispa Port Jackson (see above, and Quirk et al., 1983: 510-3).

216. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 77
‘Goose Id Bay South Coast Adiantoides crispa var’ (see above, and Quirk et al., 1983:
D10=3)):

217. Cheilanthes brown (Desv.) Domin Bennett 5(a)
‘Arnhem South Bay Point U, FebY 6 1803 2 Notholaena villea prodr 146’

Mt. Caledon, Caledon Bay, Northern Territory (Burbidge, 1956; Vallance e¢ al., in
prep.). ‘Notholaena vellea of Brown (1810: 146, entry 2‘T). Holotype (Quirk et al., 1983:
SAB).

218. Chewlanthes browni (Desv.) Domin Bennett 5(b)
‘Shoalwater Bay Conical Pine Hill’ (Perhaps ‘Notholaena vellea of Brown, 1810: 146,
entry 2-T).

219. Cheilanthes caudata R.Br. Bennett 78
‘Pteris [struck through] Cheilanthes caudata prodr 156 Port II’ Port Clinton, Queens-
land (Burbidge, 1956). Visited by Brown in August 1802 (Vallance ef al., in prep.).
Brown (1810: 156, entry 2-T). Holotype (Quirk et al. 1983: 540-1).

220. Cheilanthes distans (R.Br.) Mett. Bennett 4(a)
‘Adiantoides palacea Port Jackson |. . .] Notholaena distans prodr 146’ (“Notholaena
distans’ of Brown, 1810: 146, entry 1-J). Holotype (Quirk et a/., 1983: 526-9).

221. Chewlanthes lasiophylla Pichi-Serm. Bennett 4b
‘Adiantoides paleacea Inlet XII [. . .] Notholaena cfr N. distans’
At the Head of Spencer Gulf (Burbidge 1956), but Brown (1810: 146, entry 1) indicates

PROG. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 89

Port Jackson as the source of the material. The Prodromus entry should have the ‘M’
annotation if Spencer Gulf is meant. See Quirk et al. (1983: 531-2) and Pichi-Sermolli
(1951) regarding relationship with ‘Notholaena canescens.

222. Cheilanthes nudiuscula (R.Br.) T. Moore Bennett 60
‘Pteris denudata v[ar] N Coast Id. y2’

Pibassoo’s Island, Northern Territory (Burbidge, 1956). ‘Pteris nudiuscula of Brown
(1810: 155, entry 8-T). Holotype (Quirk e¢ al. , 1983: 537-40).

223. Cheilanthes tenuissima Bailey Bennett 61
‘Adiantoides pubescens North Coast Island y2 &C’. Pibassoo’s Island, as above.

224. Cheilanthes tenuissima Bailey Bennett 3
‘Acrostichum gracile [struck through] pteroides prodr 145 North Coast Islands 1803 RB’
(Brown 1810: 145, entry 3-IT). Type of ‘Acrostichum pteroides’ R.Br. (Quirk et al., 1983:
Saco)

225. Cheilanthes tenuifolia (Burm.f.) Sw. Bennett 72
(Syn. ssp. queenslandica Domin., see remarks on relationship of C. austrotenuifolia and C.
tenurfolia by Quirk et al. (1983: 506-13). ‘Adiantoides crispa var. North Coast Islands’
(Brown, 1810: 155, entry 1-T).

226. Cheilanthes sp.indet. Bennett 73
‘Adiantoides crispa Port II’, Port Clinton area, Queensland (Burbidge, 1956).

227. Cheilanthes sp.indet. Bennett 75
‘Adiantoides cicutaria Pteris [sic] Port IT’

228. Christella dentata (Forsk.) Browns. & Jermy Bennett 21(a)
‘5 Nephrodium molle Prodr 149 [reverse reads] Plenosouris Polypod: molle Jacq. Grose’
(Brown, 1810: 149, entry 5-J).

229. Christella dentata (Forsk.) Browns. & Jermy Bennett 21(b)
‘Polypodium molle Jacq adrip: Patersons Hunter William Rivers det 1804 [reverse
reads] 5 Nephrodium molle Prodr 149’ (Brown, 1810: 149, entry 5-J).

230. Ctenopterrs heterophylla (Labill.) Tindale Bennett 13
‘Polypodium arbor prope in caulis Dicksonia Derwent 9 Polypodium grammitidis prodr
147 (Polypodium grammitidis’ of Brown, 1810: 147, entry 9-D).

231. Culcita dubia (R.Br.) Maxon Bennett 86
Two labels: 1st reads ‘Cfr D. cicutarian [struck through] Banks of Hunter’s River Nov
1804 4 Davallia dubia prodr 157’; 2nd reads ‘Davallia dubia Port Phillip’ (Brown, 1810:
157, entry 4-J and D).

232. Culcita dubia (R.Br.) Maxon Bennett 87
‘4 Davallia dubia adripes fl. Patersons & Hunters Rivers Oct: Nov: 1804 (Brown, 1810:
157, entry 4-J).

233. Culcita dubia (R.Br.) Maxon Bennett 88
Two identical sheets “4 Davallia dubia Pt Jacksor (Brown, 1810: 157, entry 4-J).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

90 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

234. Culcita dubia (R.Br.) Maxon Bennett 89
‘4 Davallia multiflora [struck through] dubia Port Jackson’ (Brown, 1810: 157, entry 4-J).

235. Culcita dubia (R.Br.) Maxon Bennett 90
‘4 Davallia dubia prodr 157 Port Dalrymple Jan 1804 (Brown, 1810: 157, entry 4-D).

236. Cyathea australis (R.Br.) Domin Bennett 94
‘Cyalloides Kings Island Basses Strait’ (Alsophila australis’ of Brown, 1810: entry 1-D).
A tree fern, occurring in gullies near the sea. Lectotype (Tindale, 1956: 350).

237. Cyathea australis (R.Br.) Domin Bennett 95
‘1 Alsophila australis prodr 158’ (probably ‘Alsophila australis’ of Brown, 1810: 158, entry

1-J).

238. Davallia pyxidata Cav. Bennett 84
‘Davallia glycopoda P Jackson in rep (Brown, 1810: 156-7, entry 1-J).

239. Davallia pyxidata Cav. Bennett 85
Two sheets: Ist reads “Davallia pyxidata b Hunters River, Ash Island’; 2nd reads
‘Davallia glycopoda P Jackson in rup Grose [sic] (Brown, 1810: 156-7, entry 1-J).

240. Dennstaedtia davalliordes (R.Br.) T. Moore Bennett 93
‘Davallia tenera Banks of Patersons River Oct: 1804 Dicksonia davallioides prodr 158
(Brown, 1810: 158, entry 2-J).

241. Diucksonia antarctica Labill. Bennett 92
Two sheets: Ist reads “Dicksonia australis Kings Island’; 2nd reads ‘Dicksonia antarctica
Van Diemen [sic] Kings Island’ (second label relates to Brown, 1810: 157, entry 1-D. See
also next).

242. Diucksonia antarctica Labill. Bennett 91
Two labels: Ist reads ‘Dicksonia australis Derwent; 2nd reads ‘Dicksonia antarctica
prodr 157 Derwent’ (Brown, 1810: 157, entry 1-D).

243. Dicranopterts linearis (Burm.f.) L. Underw. Bennett 111
‘Polypodium dichotomum Shoalwater bay passage |. . .] 8 Gleichenia Hermanni prodr
161 (Brown, 1810: 161, entry 8-1).

244. Diplazium australe (R.Br.) Wakef. Unnumbered, but probably corresponds
to Bennett register entry 24

No original labels, but on back of sheet, not in Brown’s hand, ‘Nova cambria australe
Port Jackson (perhaps ‘Allantodia tenera R.Br., cf. Brown, 1810: 149, entry 2-J).

245. Doodia aspera R.Br. Bennett 36
‘Woodwardia aspera Banks of Hunters River frequent Oct-Novr 1804 Doodia aspera
prodr 151 (Brown, 1810: 151, entry 1-J).

246. Doodia caudata (Cav.) R.Br. Bennett 37
‘Woodwardia blechnoides sp5 [sic] Port Jackson [. ..] Doodia caudata prodr 151’
(Brown, 1810: 151, entry 3-J).

PROG. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 91

247. Doodia caudata (Cav.) R.Br. Bennett 38
‘Woodwardia Port Dalrymple Jan: 1804 (Brown, 1810: 151, entry 3-D).

248. Doryopterts concolor (Langsd. & Fisch.) Kuhn Bennett 62
‘Pteris pedata prodr 155 Broad Sound in Montibus prope Upper Head’ (Brown, 1810:
155), cain O30).

249. Drynaria rigidula (Sw.) Bedd. Bennett 14a (but should be 15)
‘Polypodium anomalum Carpentaria Groote Eyland Frondes sterilis sumatae breviores
latiores [reverse reads] Polypodium quercifolium B non Lin’ (Brown, 1810: 147, entry

10-1).

250. Drynaria sparsisora (Desv.)'T. Moore Bennett 14b (bis)
10 Polypodium quercifolium Prodr 147 Carpentaria Coen River (Brown, 1810: 147,
entry 10-T).

251. Drynaria sparsisora (Desv.) T. Moore Bennett 14b (bis)
‘Polypodium Keppel Bay (Brown, 1810: 147, entry 10-T).

252. Gleichenia alpina R.Br. Bennett 105
‘3 Gleichenia alpina desc prodr 160 Summitat Table Mountain Derwent’ (Brown, 1810:
161, entry 3-D).

253. Gleichenia dicarpa R.Br. Bennett 108
(Syns. ‘Gleichenia circinnata’ Sw., ‘Calymella circinnata (Sw.) Ching.?)

‘Gleichenia pubescens sp [. . .] P Dalrymple desc’ (possibly Brown, 1810: 161, entry
5-D).

254. Gleichenia microphylla R.Br. Bennett 106
(Syn. ‘Calymella circinnata’ (Sw.) Ching.?)
‘Gleichenia cfr speluncae Port Jackson’ (Brown, 1810: 161, entry 4-J).

255. Gleichenia microphylla R.Br. Bennett 107
Two labels: 1st reads “4 Gleichenia microphylla Prodr 161 Kings Island’; 2nd reads ‘Port
Jackson’ (Brown, 1810: 161, entry 4-.D (label 1) and J (label 2) ).

256. Gleichenia microphylla R.Br. Unnumbered
No original label: secondary label reads ‘New Holland Kents Group Dec 1803’ (Brown,
1810: 161, entry 4-D).

257. Gleichenia rupestris R.Br. Bennett 104
(Syn. ‘Calymella rupestris’ (R.Br.) Ching.)

‘Gleichenia austriaca desc Port Jackson [. . .] 2 Gleichenia rupestris Port Jackson prodr
160’ (Brown, 1810: 160, entry 2-J).

258. Gleichenia speluncae R.Br. Bennett 103
(Syn. ‘G. circinnata var.?, or ‘G. microphylla R.Br. °).
‘Gleichenia subsimplex desc rup [. . .| Port Jackson’ (Brown, 1810: 160, entry 1-J).

PROG. LINN. SOC. N.S.W., 111 (2), 1989

92 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

259. Grammitis billardierr Willd. Bennett 6b &c
Label 6b reads ‘Ceterach polypodiodes rupestris in later Mont Tabul Derwent Gram-
mitis australis prodr 146’ (Brown, 1810: 146, entry 1-D); 6c reads ‘Grammitis australis
prodr 146 Port Jackson 1804 RB’ (Brown, 1810: 146, entry 1-J).

260. Histiopterts incisa (Thunb.) J.Sm. Bennett 52
‘3 Pteris vespertilionis P Jack[son and] Derwent (Brown, 1810: 154, entry 3-J).

261. Hustiopteris incisa (Thunb.) J.Sm. Bennett 54
‘3 Pteris vespertilonis prodr 154 Kings Island’ (Brown, 1810: 154, entry 3-D).

262. Histiopterrs incisa (Thunb.) J.Sm. Bennett 55
‘Pteris vespertilonis B Port Jackson’ (Brown, 1810: 154, entry 3-J).

263. Hymenophyllum cupressiforme Labill. Bennett 100
‘Hymenophyllum tunbridgense prodr 159 Derwent’ (Brown, 1810: 159, entry 4-D).

264. Hymenophyllum flabellatum Labill. Bennett 98
‘Hymenophyllum nitens prodr 159 Derwent (Brown, 1810: 159, entry 2-D).

265. Hypolepis rugosula (Labill.) J.Sm. Bennett 16a
‘12 Polypodium rugosulum Prod 147 [reverse reads] Polypodium affine Kings Island’
(Brown, 1810: 147, entry 12-D).

266. Lastreopsis decomposita (R.Br.) Tindale Bennett 22
‘Pteridioides acuminata Polypodium Involuc renif. adrip. Hunters River Oct 1804 7
Nephrodium decompositum prodr 149’.

(‘Nephrodium decompositum’ of Brown, 1810: 149, entry 6-J). Holotype of L. decomposita
(cf. Tindale, 1965: 320-3).

267. Lastreopsis decomposita (R.Br.) Tindale Bennett 23
‘Polypodium tenerum Broad Sound Peaked West Hill in sylvis umbrosis [. . .] 7 Nephro-
dium decompositum [struck through] tenerum prodr 149’ (Nephrodium tenerum’ of
Brown, 1810: 149, entry 7-T).

268. Lindsaea ensifolia Sw. Bennett 80
‘Lindsaea lanceolata Prodr 156 [reverse reads] Lindsaea pleroides [‘Vittaria struck
through] Port II Shoal Water Bay passage’ (Brown, 1810: 156, entry 2-T). This specimen
has been determined by K. U. Kramer in 1965 as the ssp. agatz (Brack.) Kramer.

269. Lindsaea ensifolia Sw. Bennett 81
‘2 Lindsaea lanceolata [@] [reverse reads] Lindsaea polymorpha North Coast Island Y,’
Arnhem Bay area, N.T. (Burbidge, 1956; Brown, 1810: 156, entry 2-T). Determined by
K. U. Kramer in 1965 as the ssp. agai (Brack.) Kramer.

270. Lindsaea linearis Sw. Bennett 79
Two labels: 1st reads ‘Lindsaea recurifera Port Jackson [reverse reads] Lindsaea linearis
Prodr 156’; 2nd reads ‘1 Lindsaea linearis’ (Brown, 1810: 156, entry 1-J, and Kramer and
Tindale, 1976: 112-3).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 93

271. Lindsaea microphylla Sw. Bennett 83
Original label damaged, remaining part reads ‘[. . .]| Port Jackson’ (Brown, 1810: 156,
entry 4-J, and Kramer and Tindale, 1976: 95-8).

272. Lindsaea media R.Br. Bennett 82
‘Lindsaea irregularis North Coast Island y2 Feb 18-24 1803 Z Pibassoo’s Island
(Burbidge, 1956). (Brown, 1810: 156, entry 3-I, and Kramer and Tindale, 1976: 98-101).

273. Lygodium flexuosum (L.) Sw. Unnumbered but probably corresponds

to Bennett register entry 116
‘Nova Hollandia or septen trionalis 1803 RB’ (Lygodium semibipinnatum’ of Brown,
oO G2 entry 2-1 ):

274. Lygodium microphyllum (Cav.) R.Br. Unnumbered but probably corresponds
(Syn. ‘Lygodium scandens (L.) Sw.) to Bennett register entry 115
No original label; secondary labels reads ‘Nova Cambria australis inter tropicum RB’
(Lygodium microphyllum’ of Brown, 1810: 162, entry 1-T).

275. Marsilea angustifolia R.Br. Bennett 141
‘3. Marsilea angustifolium Carpentaria Island g Dec 25 1802’, Vanderlins Island
(Burbidge, 1956; cf. Vallance et al., in prep.). (Brown, 1810: 167, entry 3-T).

276. Marsilea hirsuta R.Br. Bennett 138
‘Marsilea hirsuta Carpentaria Island f, Bountiful Island (Burbidge, 1956). Brown was
there on 3 December 1802 (Vallance et al. in prep.). (Brown, 1810: 167, entry 2-T).

277. Marsilea hirsuta R.Br. Bennett 139
‘2 Marsilea [quadrifolia struck through] hirsuta Broad Sound in frales de propes scites
temporebus inundatis (perhaps Brown, 1810: 167, entry 2-T). ‘Marsilia quadrifolia’ of
Brown's Diary, collected on 15 September 1802 (Vallance et a/., in prep.).

278. Marsilea hirsuta R.Br. Bennett 140
‘2 Marsilea villosa [struck through] hirsuta in |. . .] Port Jackson’ (Brown, 1810: 167,
entry 2-]).

279. Marsilea mutica Mett. Bennett 137
(Syn. ‘M. brown’ R.Br., cf. Beadle e¢ al. (1972), in index.)
‘1 Marsilea quadrifolia Port Jackson’ (Brown, 1810: 167, entry 1-J).

280. Marsilea mutica Mett. Unnumbered
Two labels: 1st reads ‘Nova Cambria australis Port Jackson R Br’; 2nd reads ‘Marsilia
australis RB Marsilia quadrifolia prodr [blank] (probably Brown, 1810: 167, entry 1-J).

281. Mecodium australe (Willd.) Copel Bennett 99
‘3 Hymenophyllum flabellatum [‘undulatum’ struck through] Labill prodr 159 Derwent’
(Hymenophyllum flabellatum’ of Brown, 1810: 159, entry 3-D).

282. Mecodium rarum (R.Br.) Copel. Bennett 97
Two labels: Ist reads ‘1 Hymenophyllum rarum prodr 159 Derwent; 2nd _ reads
‘Hymenophyllum rarum prodr |. . .] Van Diemans Island 1804 RB’ (Brown, 1810: 159,
entry 1-D).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

94 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

283. Maicrosorum punctatum (L.) Copel. Bennett 10
‘5 Polypodium irioides [‘punctatum’ struck through] Prodr 147 Keppel Bay Shoal Water
Bay Broad Sound’. Another sheet of the same gathering has a secondary label which
reads ‘Nova Cambria Australis intra tropicum 1802 RB’ (Brown, 1810: 147, entry 5-I).

284. Nephrolepis hirsutula (Forst.f.) C. Presl Bennett 20
‘Nephrodium exaltatum prodr 148 Plendioides davilliastrum desc [“Davallioides’ struck
through] Port II East Coast’. Port Clinton, Queensland (Burbidge, 1956), (Brown, 1810:
148, entry 2-T).

285. Ophioglossum costatum R.Br. | Bennett 118
‘2 Ophioglossum costatum Arnhem N Bay Feb 14 1803 desc 55’ (Brown, 1810: 163, entry
ZAM,

286. FPellaea falcata (R.Br.) Fee Bennett 59
‘Kents Islands & Hawkesbury 6 Pteris [‘adiantoides’ struck through] falcata prodr 15¥
(‘Pteris falcata of Brown, 1810: 154, entry 6-J and D).

287. FPellaea paradoxa (R.Br.) Hook. Bennett 63
‘Port Jackson 1 Adiantum paradoxum prodr 155’ (‘Adiantum paradoxum’ of Brown,
1810: 155, entry 1-J).

288. Phymatosorus diversifolia (Willd.) Pichi Serm. Bennett lla-c
Three sheets: 1st (Bennett Ila) reads ‘6 Polypodium billarderi prodr 147 Port Dalrymple
in umbrosis & truncus arbor Jan 1804’; 2nd reads (Bennett 11b) “‘Polypodium poly-
morphum var Port Jackson’; 3rd (Bennett 11c) reads ‘Kents Group Bass Strait 7 Poly-
podium [‘lycopodioides’ struck through] Billardieri prodr 147’ ((Polypodium billardiery
of Brown, 1810: 147, entry 7-D). The indication would appear to be that the Port
Jackson specimen did not feature in the description.

289. Platycerrum bifurcatum (Cavy.) C.Chr. Bennett 1
Two labels: Ist reads ‘Acrostichum alcicorne prodr pl45 Port Jackson 1802-5 Rocks &
‘Trees’; 2nd reads ‘1 Acrostichum alcicorne prodr 145’ (Brown, 1810: 145, entry 1-J).

290. Platyzoma microphyllum R.Br. Bennett 101
(?Syn. ‘Gleichenia platyzoma’ F. Muell.)

‘Platyzoma microphylla Carpentaria Island h’ (North Island cf. Burbidge 1956) (Brown,
1810: 160, entry 1-T).

291. Platyzoma microphyllum R.Br. Bennett 102
(? Syn. ‘G. platyzoma F. Muell.)

‘Acrostichum? Gleichenia? Facing Island Port I’ (Queensland, cf. Brown, 1810: 160,
entry 1-I).

292. Pleurosorus rutiflorus (R.Br.) Fee Bennett 7
‘Derwent Risdon 2 Grammitis rutaefolia prodr 146’. Another specimen on this sheet
bears the secondary locality ‘Nova Hollandia ora australis Kangaroo Island 1802 RB’
(probably ‘Grammitis rutaefolia’ of Brown, 1810: 146, entry 2-D).

PROC. LINN. SOC. N.S.W,, 111 (2), 1989

E. W. GROVES AND D. T. MOORE 5

293. Polyphlebrum venosum (R.Br.) Copel. Bennett 96
‘Derwent 1 Trichomanes [‘repandum’ struck through] venosum prodr 159’ (Brown, 1810:
159, entry 1-D).

294. Polystichum australiense Vindale Bennett 18
No original label; secondary annotation reads ‘Derwent’. Separated out from material of
Bennett 17 (see below) and assigned by Bennett’s workers to No 18.

295. FPolystichum proliferum (R.Br.) C. Presl Bennett 17
‘Aspidium proliferum prodr 147 desc Port Jackson [and] Derwent’ (Brown 1810: 147-8,
entry 1-D). Lectotype (Tindale, 1961: 50).

296. Pteridium esculentum (Forst.f.) Cockayne Bennett 57
‘Pteris esculenta var P Jackson’ (Brown, 1810: 154, entry 5-J).

297. Pteridium esculentum (Forst.f.) Cockayne Bennett 58
‘D Pteris esculenta prodr 154 Derwent (Brown, 1810: 154, entry 5-D).

298. Pteris tremula R.Br. Bennett 53
‘Grose’ (Brown, 1810: 154, entry 4-J).

299. Pteris tremula R.Br. Unnumbered but corresponds to Bennett register entry 56
‘Pteris tremula prodr 154 Port Jackson’ (Brown, 1810: 154, entry 4-J).

300. Pteris umbrosa R.Br. Bennett 51
‘Pteris umbrosa First branch of the Grose Pteris’ (Brown, 1810: 154, entry 1-J).

301. Pyrrosia confluens (R.Br.) Ching Bennett 9
‘Pteroides varia [struck through] Polypodium confluens prodr 146 cfr Plenopilo-
selloides ad rip Hunters & Patersons Rivers Oct: Nov 180¥ (‘Polypodium confluens’ of
Brown, 1810: 146, entry 3-J).

302. Pyrrosia rupestris (R.Br.) Ching Bennett 8
‘Polypodium rupestre prodr 146 [reverse] Polypodium serpens B & S Forst Prod: Vahl
Symb Port Jackson’ (‘Polypodium rupestris’ of Brown, 1810: 146, entry 2-J).

303. Rumohra adiantiformis (Forst.f.) Ching Bennett 19
‘2 Aspidium coriaceum prodr 148 Polypodium adiantiforme Forst prod Derwent adras
montis (‘Aspidium coriaceum of Brown, 1810: 148, entry 2-D).

304. Schizaea bifida Willd. Bennett 113
Two labels: 1st reads; ‘3 Schizaea furcata [struck through] bifida prodr 162 Port II’; 2nd
reads ‘Schizaea furcata Richmond’ (Brown, 1810: 162, entry 3-J. The provenance
annotation is J and D; perhaps T and J are meant).

305. Schizaea bifida Willd. Unnumbered — Dryander duplicate, see introduction.
‘Acrostichum pectinastordes Nova Cambria australis Port Jackson’.

306. Schizaea dichotoma (L.) J.Sm. Bennett 114
‘Schizaea dichotoma 4 Schizaea aspera scabr cfr S dichotom: prodr 162’ (Brown, 1810:
162, entry 4-T).

PROG. LINN. SOG. N.S.W,, 111 (2), 1989

96 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

307. Schizaea fistulosa Labill. Unnumbered
“Terra Diemam (‘Schizaea fistulosa of Brown, 1810: 162, entry 2-D).

308. Schizaea fistulosa Labill. Unnumbered
(Syn. ‘Microschizaea fistulosa’ (Labill.) Reed).

‘Schizaea fistula[. . .]. Locality probably Tasmania as above (cf. Brown, 1810: 162, entry
eID),

309. Schizaea rupestris R.Br. Bennett 112
‘Schizaea rupestris prodr 162 desc 25 nep |. . .| Port Jackson’ (Brown, 1810: 162, entry
1-J).

310. Stucherus flabellatus (R.Br.) St. John Bennett 109
‘Port Jackson 6 Gleichenia [‘prolifer’ struck through] flabellata prodr 161 (‘Gleichenia
flabellata of Brown, 1810: 161, entry 6-J).

311. Sticherus tener (R.Br.) Ching Bennett 110
‘7 Gleichenia tenera prodr 161 in rup. hum ad rip rivul alpin Mon|tis] Tabul{aris]’
(‘Gleichenia tenera of Brown, 1810: 161, entry 7-D).

312. ‘odea barbara (L.) T. Moore Bennett 117
‘Osmunda barbara prodr 163 Port Jackson’ (Brown, 1810: 163, entry 1-J).

313. Vittaria elongata Sw. Bennett 50
‘Vittaria acostata Broad Sound Peaked West Hill parasit: super Asplen|ium] nidus’
(Brown, 1810: 153, entry 1-T).

CYCADOPHYTES
Cycadaceae

314. Cycas angulata R.Br. Bennett 3106 (pro parte)
No original label. According to an annotation by J. R. Maconochie of 18 November
1976, the specimen is probably from Bountiful Island, Gulf of Carpentaria, and the “I”
entry confirms the tropical provenance (Brown, 1810: 348, entry 2-T). In his Diary
(Vallance et al., in prep.) Brown does not mention a cycad being collected from Bountiful
Island but Good mentions a ‘Cycas circinalis’ which his editor (Edwards, 1981: 104)
identified as C. media R.Br. But we believe that C. angulata is correct. Bauer painting
159/60, Botany Library, BM (NH), is probably based on this plant.

315. Cycas media R.Br. Bennett 3106 (pro parte)
No original label. According to an annotation by J. R. Maconochie of 18 November
1976, the specimen is probably from Cumberland Islands, Queensland. The “IT” entry
(Brown, 1810: 348, entry 1-I) confirms this (cf. Mabberley, 1985: plate 4). Certainly
Brown collected a specimen of ‘Cycas circinalis’ at Calder Island (Cumberland Islands)
on 16 October 1802 which could be this specimen. BM (NH) Bauer painting 157/8 is
probably based on this plant (Vallance e¢ al., in prep.).

CONIFEROPHYTES
Araucariaceae

316. Araucaria cunninghami Ait. Bennett 3115

PROG. LINN. SOG. N.S.W.,, 111 (2), 1989

E. W. GROVES AND D. T. MOORE 97

‘Araucaria cunninghamii Port Bowen 1802 RB’ (probably the specimen collected 21
August 1802 according to Brown’s Diary (Vallance et al., in prep).

Cupressaceae

317. Callitris gracilis R. Baker Bennett 3108
(Syn. ‘Frenela robusta’ A. Cunn.)
Two labels: 1st reads ‘Callitris glauca Nob Mount Brown Spencers Gulf 1802 RB’; 2nd
reads ‘Genus Conifer Inlet XII South C’. Gathered from Mount Brown, 10-11 March
1802 (cf. Vallance et a/., in prep.).

318. Callitris gracilis R. Baker Bennett 3109
(Syns ‘Frenela robusta’ A. Cunn., ‘Callitris propinqua’ R.Br.)

‘Callitris propinqua Nob Kanguroo Island 1802 RB’ Gathered March 1802 (cf. Vallance
et al., in prep.).

319. Callitris muellert (Parl.) F. Muell. Bennett 3112
(Syn. ‘Frenela mueller’ Parl., cf. Garden, 1956: 383).
‘Callitris Nova Holland loc incerta.

320. Callitris rhomboidea R.Br. ex A. & L. C. Rich Bennett 3107
(Syn. Holotype of ‘Callitris ventenati’ Mirb., Garden, 1956: 386)
‘Callitris ventenati1 Nob Port Jackson 1804-5”

321. Callitris rhomboidea R.Br. ex A. & L. C. Rich Bennett 3114
(Syn. ‘Callitris australis’, cf. Garden, 1956: 386).

‘Genus Coniferar: adripas saxosus fluvii Cataract River Port Dalrymple Jan 10 1804
[. . .] Callitris australis’

322. Callitris tuberculata R.Br. ex R. T. Baker & H. G. Smith Bennett 3110
(Syn. ‘Frenela robusta var. verrucosa, also ‘Callitris preiss1’ Miq.).

Two labels: Ist reads ‘Middle Island of Goose Island Bay 1802 RB’; 2nd reads ‘Genus
Conif Bay II South Coast 1802 Jan 12’. Gathered January 1802 from the Archipelago of
the Recherche (cf. Willis, 1959: 99; and Vallance et al., in prep.).

323. Callitris sp.indet. Bennett 3111
‘Port Jackson.
324. Callitris columellaris F. Muell. Bennett 3113

‘Nov gen Conifer Sandy Cape Arbor Mediocris cortice ramoso crasso’. Collected 30 July
1802, Vallance et al. (in prep.).

Podocarpaceae

325. Lagarostrobos franklini (Hook.t.) C. J. Quinn Bennett 3120
‘Huon Pine’. No locality given on original label. This taxon sensu Quinn (1982).

326. Phyllocladus aspleniifolius (Labill.) Hook.f. Bennett 3119
(Syn. ‘Podocarpus asplenifolia’ Labill.).
‘Derwent [. . .] Podocarpus asplenifolia.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

98 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

327. Podocarpus elatus R.Br. ex Endl. Bennett 3117
Two labels: Ist reads “Taxus cfr T elongatum Hunters River; 2nd reads “Iaxus cfr T
elongatum In sylvis prope ripas fluvio Paterson A A Ash Island Insulas que adjacent
Cabbage tree swamp & C Desc In mscr N C No95’. Probably gathered 14 October 1804
(cf. Vallance et al. in prep.).

328. Podocarpus lawrence: Hook.f. Bennett 3116
(‘Podocarpus alpinus’ of de Laubenfels, 1985).

“Taxus emersa Nol20 desc Misc fl Derwent In Summitate saxosa Montis Tabularis
prope fluio: Derwent Apr: Mai 180.

329. Podocarpus spinulosus (Sm.) R.Br. ex Mirb. Bennett 3118
No original label.

INDEX

Poorly-localized material is unindexed, but material from the south coast is given
below. This is broadly equivalent to Brown’s (1810) annotation ‘M’:
3, PO, 4), 13, 20, 22, Bae, DD, BO=T, 42-3), 42], DD 13), GH GO, IZZ, WA) 72, UTS), Zil4e, SUS
and 322.

Material from Tasmania, and Bass Strait, approximately equivalent to Brown’s
(1810) ‘D’ annotation:
Dy 4, Dy Il, WIG), Zl, ZH, HlleB2,. HAs, DS, Gel, 43, HS=Hil, DS), DY, OA, GIS, /le3, 13-9, O7/, OY.
GHG, 100, WOSabL, NSD, Wl7/3, 20225), IOS, W378, Wal, Wa 3=59, ID /=3,, IOW04, loo=7,
WOME, W7/H,, USV-GO. Ws, AOI-O42, AVO=bl, ZHO=l, AIH=O, Z4wle4o2,, Wei), VU=3, YO, ZO, VOS=
5, ZMleZ, ZO, 2HS, 22-3, 299, 27, 303, 307-8, Dbl, BAN, BAG eral HAS:

Material from New South Wales, equivalent to Brown’s (1810) ‘J’ annotation:
QI-D, WOIO4, WG-29, UBS, WA), We, WHO, HOS, Wers,, W7Sie7/4s, WO, MOA, WSHecks, I=},
IQG-20Z, AVS, AIDA, VAY, ZAS=2Y), VBL=S4e, ZDI-B), VAL), WEA, VIO), BOL, AEG, Z/0=
Mi, 2TS-80, 237, ZE9, 296, ZVS-3OZ, SO, SVQ, Si2, 320, S23 aval S27.

Tropical material, equivalent to Brown’s (1810) “TI” annotation:
Di), 29), 48, BO), 82, 84, W/T=78, MSS, DD, ZiT, LAA, ZHAaVS), 243}, ZAG-Gil, VOT, AOS=59),
DIDI Ny POSBO, 2YO-DL, BOL, BOO, BBall, BLA? aanel B27.

INDEX OF BENNETT NUMBERS vs GROVES & MOORE NUMBERS
Wineee aie In lskernneitt mvtonl serge (3, Sa, Bis, 440), HO), Ges, GY, IG, WS), iil, ISS (| =25)]).
ISO SHO), WN Sail, i225], Io [S245], 200 ||P Seer], Zl) ease ares |p 24], 256,
273 (2? =16), 274 [2=155], 280, 299 |=56], 305, 307 and 308. Possibly equivalent
Bennett numbers are given in brackets — see text for each such entry. For Groves &
Moore numbers marked with an asterisk (*) there is no Bennett number.

Bennett Groves & Moore Bennett Groves& Moore Bennett Groves & Moore

1 289 9 301 19 508

2 a 10 283 20 284

3) 224 lla-c 288 2la 228

4a 220 12 186 21b 229

4b Pa 13 230 BD 266

al 27, allo) 249 BS 267

5b 218 14b BIO) Ce LHL 24 244*

6b &c 259 l6a 265 25 188, 192 & 193*
7 292 17 295 Di 194

8 302 18 294 28 196*

PROG. LINN. SOC. N.S.W., 111 (2), 1989

E. W. GROVES AND D. T. MOORE 99

Bennett Groves & Moore Bennett Groves & Moore Bennett Groves & Moore

29 Is 81 269 131 168
30 tO Or 82 272 132 173
31 Oe 83 ZH 133 172
33 187 84 238 13)4¢ 197
35 189 85 L39) 139 198
36 245 86 POI 136 199
on 246 87 23y2 137 DUS
38 247 88 D9) 138 276
39 202 89 Dove 139 ZU
40 205 90 235 140 278
41 201 Dil 242 141 ZAS
42 203 92 Deel 157 39
43 204 93 240 158 41
44 209 & 210* 94 236 159 aes
45 207 95 ZU 161 44
46 208 96 2S)3} 162 45
ALT 206* a 282 163 46
48 211 & 212 98 264 166 3
49 200 yg 281 167 5
50 ols 100 263 168 6
51 300 101 290 169 7
oy 260 102 291 170 10
53 298 103 258 171 9)
54 261 104 257 UZ 4
55 262 105 DDD 173 2
56 DS ie 106 DO: 174 11
5/) 296 107 Dd)5) 7 32
58 ZS)i/ 108 ZS 176 69
a9) 286 109 310 177 a)
60 222 110 311 179 18
61 LS 111 243 181 2, ds} we wee
62 248 112 309 18la 15
63 287 113 304 182 a2
64 183 114 306 193 60
65 184 115? Dae 195 53
66 185 116? Duss 203 16
67 181 117 312 205 1
68 182 118 285 206 68
69 178 119 213 208 20
70 7) 120 174 209 22
ML 180 121 176 210 42
Ue 229 122 175 211 2
73 226 123 169 PS 7
74 Dist 124 170 214 21
75 2h 125 171 215 48
76 215 126 163 216 47
el 216 127 164 217 oy
78 ZAG 128 165 218 27
19 270 129 166 219 Uae
80 268 130 167 221 25

PROG. LINN. SOC. N.S.W., 111 (2), 1989

100 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS

Bennett Groves & Moore Bennett Groves & Moore Bennett Groves & Moore

923 28 984 150 539 100
994 26 985 157 540 116 & 119
995 49 986 160 542 149

226 19 287 155 543 111

BAT 3D) 288 oul 544 128

228 50 289 152 545 134

229 31 290 144 546 97
ie Sr t Soecie
933 78 993 143 i ae
235 74 294 ie a iz

936 63 995 14 See!
240 54 296 149 eee eee
244 16 501 129 I! tee

245 55 502 124 399 133

249 li 504 99 556 112

250 57 507) 104 558 137

951 30 & 71 511 93 560 102

952 72 515 107 961 101

253 64 516 118 963 96

254 61 Ry 130 565 109

957 65 518 139 & 140 566 110

959 62 519 126 567 120

260 67 520 136 569 103

261 80 521 114 3106 314 & 315
963 66 522 98 3107 320

264 56 HLS 138 3108 7

266 75 S28 131 3109 318

267 79 525 141 3110 399

268 73 526 105 & 132 3111 393

969 59 527 91 3119 319

971 51 528 135 ove AO.

972 58 530 90 tnd <

276 83 531 89 i

277 81 532 85 ce Ve

279 82 533 86 3116 328

280 162 534 87 & 88 S117 327

281 155 536 115 3118 329

282 154 537 113 3119 326

983 158 538 117 3120 395

ACKNOWLEDGEMENTS

A work of this nature could not have been undertaken without the assistance of
specialist taxonomists and curators. Consequently in the British Museum (Natural
History) we wish to record our thanks to Miss J. M. Camus, Dr D. J. Galloway, Mr L.
Ellis, Dr A. J. Harrington, Dr C. R. Hill, Mr S. I. Honey, Mrs L. M. Irvine, Mr J. H.
Price, and Mr I. Tittley. We also thank the Botany Librarian, Mrs C. V. Ellwood, for
access to the Brown slip catalogue and Mr J. F. M. Cannon, Keeper of Botany, also for
access to the collections in his charge.

We owe a considerable debt to our Australian friends, particularly to Mr K. Hill

PROG. LINN. SOG. N.S.W., 111 (2), 1989

E. W. GROVES AND D. ‘T. MOORE 101

and Mrs Karen Wilson of the Royal Botanic Gardens, Sydney, and also Professor T. G.
Vallance of the University of Sydney. We thank Dr I. R. Price, James Cook University,
Queensland, for information on Brown’s algae, and Dr M. A. Clements of the
Australian National Botanic Gardens, Canberra, for helpful discussion.

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PROC. LINN. SOC. N.S.W., 111 (2), 1989

The Lichens of Norfolk Island

1: Introduction and the Family Parmeliaceae

JOHN A. ELIX and HEINAR STREIMANN

ELIx, J. A., & STREIMANN, H. The lichens of Norfolk Island. 1: Introduction and the
family Parmeliaceac. Proc. Linn. Soc. N.S.W. 111 (2), 1989: 103-121.

The historical records of lichens of the Norfolk Island complex are discussed and
synonymy given. The 16 species (from 7 genera) of the Parmeliaceae which currently
grow there are examined critically. Each species is described fully (including chemistry)
and its distribution is recorded. The endemic Flavoparmelia norfolkensis Elix & Streim. is
described as new to science. All 16 species on these geologically recent, oceanic islands
possessed vegetative dispores, attesting to the efficiency of this method of reproduction.
John A. Elix, Department of Chemistry, The Faculties, Australian National University, G.P-O. Box
4, Canberra, Australia 2601, and Heinar Streimann, Cryptogamic Herbarium, Australian National
Botanic Gardens, G.P.O. Box 1777, Canberra, Australia 2601; manuscript receved 23 November
1988, accepted for publication 15 February 1989.

KEY WORDS Flavoparmelia norfolkensis Elix & Streim., Neofuscelia verrucella (Essl.) Essl.,
Paraparmelia scotophylla (Kurok.) Elix & Johnston, Parmelia erumpens Kurok., Parmelinopsis
spumosa (Asahina) Elix & Hale, Farmotrema austrocetratum Elix & Johnston, Parmotrema
chinense (Osbeck) Hale & Ahti, Parmotrema crinitum (Ach.) Choisy, Parmotrema cristiferum
(Taylor) Hale, Parmotrema gardnen (Dodge) Sérusiaux, Parmotrema rampoddense (Nyl.)
Hale, Parmotrema reticulatum (Taylor) Choisy, Parmotrema sancti-angelu (Lynge) Hale,
Parmotrema tinctorum (Despr. ex Nyl.) Hale, Xanthoparmelia amplexula (Stirton) Ehx &
Johnston, Xanthoparmelia australasica D. Gall., Parmeliaceae, chemotaxonomy, Norfolk
Island, lichens.

INTRODUCTION

The Norfolk Island complex (including Norfolk, Phillip and Nepean Islands with
the satellite islands) is an isolated volcanic outcrop in the South Pacific Ocean between
longitudes 167°55’ and 168°00’E and latitudes 28°59’ and 29°08’S. The islands lie on
a narrow, steep-sided submarine ridge (the Norfolk Ridge) which extends from New
Caledonia to New Zealand (Jones and McDougall, 1973). The closest land is New
Caledonia, 670 kilometres to the north. Norfolk Island les approximately 1610
kilometres north-east of Sydney and_1100 kilometres north-north-west of Auckland
(Eig):

The islands are an Australian territory: Norfolk Island (the largest) is 8 kilometres
long and 6 kilometres wide; Phillip Island is 2 kilometres long and 1 kilometre wide,
while Nepean Island is approximately 500 metres long and 300 metres wide (Figure 2).

Norfolk and Phillip Islands have similar geology: both are almost completely
volcanic in origin. Olivine basalt lavas predominate on Norfolk Island, with basaltic
tuffs relatively more common on Phillip Island. This volcanic activity occurred 2.3-3.2
million years ago, during the Pliocene epoch (Jones and McDougall, 1973). Nepean
Island and part of Norfolk Island (near Kingston) consist of coarse marine calcareous
rock of late Pleistocene origin (Ovington, 1984).

The highest point on Norfolk Island is Mt Bates, which is 319 metres above sea
level. Mt Pitt is slightly lower (318m), but these peaks are only 600 metres apart and
together form the summit complex. The southern half and north-west corner of the
island are gullied plateaux 90-120 metres a.s.]. and are distinct from the Mt Pitt — Mt
Bates summit area. Most of the coastline, except for a length on the southern coast

(Kingston area) and several bays, consists of cliffs up to 140m high (Jones and
McDougall, 1973; Ovington, 1984).

PROC. LINN. SOC. N.S.W., 111 (2), 1989

104 LICHENS OF NORFOLK ISLAND 1

Fig. 1. Location of Norfolk Island.

Phillip Island is even more rugged and precipitous, rising to its highest point at
Jacky Jacky (280 metres a.s.].) and surrounded by coastal cliffs. Largely defoliated by
feral rabbits, pigs and goats, the island is eroded extensively, with many exposed boul-
ders and rock outcrops. The plateau above the coastal cliffs consists of a series of eroded
valleys which rise towards the summit, where the slope ends abruptly at a near-vertical
cliff, 280 metres a.s.l. (Coyne, 1982). Nepean Island, an uninhabited limestone island a
few acres in extent, rises to 32 metres (Hoare, 1965).

The climate of Norfolk Island is sub-tropical with no extremes of temperature and
moderate relative humidity (72-81%) with no seasonal pattern. Mean monthly tem-
perature fluctuates from minima near 12°C (in winter) to maxima near 25°C (in
summer) and the daily range is rarely more than 8°C. Rainfall (mean annual precipita-
tion 1313mm) occurs throughout the year with a winter maximum (June average of
163mm). Droughts of some weeks duration may occur (Ovington, 1984).

HISTORY OF LICHENOLOGICAL INVESTIGATIONS

Captain James Cook discovered Norfolk Island on October 10, 1774, during his
second voyage around the world. The following day he landed at Duncombe Bay and
named the island after the then Duchess of Norfolk. Although uninhabited at the time,
Polynesian stone adzes have been discovered since, suggesting earlier settlement. Four-
teen years after discovery, the first of two penal settlements was established and main-
tained until 1814. A second penal settlement was established in 1826 and abandoned 30
years later. In 1856, descendants of the ‘Bounty’ mutineers left their home on Pitcairn
Island and established the first free settlement on Norfolk, and today the ‘Islanders’ still
comprise some 30% of the total population of approximately 2,000 (Hoare, 1965).

Norfolk Island has been visited by a number of scientific expeditions in the course
of its short history. Few of the early expeditions contributed greatly to the lichen flora.

PROG. LINN. SOG. N.SW., 111 (2), 1989

J. A. ELIX AND H. STREIMANN 105

Captain Cook Mem.

A Mt Bates NORFOLK ISLAND

A Mt Pitt

Cascade Ba
0} Flat Rock X Broken Pine y

X "The Cockpit"

Ball Bay

Pt. Hunter

4 NEPEAN ISLAND

PHILIP ISLAND

167°5S'

Fig. 2. Map of the Norfolk Island complex. Here, and on Fig. 3, ‘Philip Island’ should be Phillip Island.

In 1804-5 Ferdinand Bauer visited the island and his collections and drawings were
submitted to Stephan Endlicher of Vienna, who published a Prodromus (Endlicher, 1833
to the plants which listed four lichens: Evernia (Usnea) melaxantha Ach., Parmelia (Parmo-
trema) perlata Ach., Parmelia (Physcia) caesia Ach., and Sticta (Pseudocyphellaria) aurata Ach.

In 1830 Allan Cunningham visited the island and augmented the knowledge of its

PROG. LINN. SOG. N.S.W., 111 (2), 1989

106 LICHENS OF NORFOLK ISLAND 1

botany; his notes were published posthumously in Heward (1842). A lichen not enumer-
ated by Endlicher was listed, namely Ramalina scopulorum Ach.

Baron Ferdinand von Mueller subsequently investigated the botany of the island on
the basis of specimens collected by Isaac Robinson, a resident and agent for the Sydney
Botanic Gardens. The identity of the lichen collections was communicated subsequently
to J. H. Maiden (see below).

In 1904 J. H. Maiden (Director of the Botanic Gardens in Sydney) reported a total
of 29 lichen species for Norfolk Island. This list was prepared by Edwin Cheel (Botanic
Gardens, Sydney) after examining the collections of Maiden and Boorman (made
during a visit to the island in 1902), and was supplemented by F. von Mueller (Mel-
bourne), who supplied the names of six species identified by J. Muller of Aargau,
Switzerland.

This list is presented here using currently-accepted names for taxa (bold) followed,
where appropriate, by the synonyms used by Maiden in brackets. [?] infers a dubious
identification — according to Maiden. The species list included: Anaptychia ciliaris (L.)
Korb. [?]; Catinaria grossa (Pers. ex Nyl.) Vainio {Patellaria grossa Mull. Arg.}; Catinaria
versicolor (Fée) Sipman {Patellaria versicolor Fée}; Chiodecton perplexum Nyl.; Clathro-
porina eminentior Nyl.; Coenogonium implexum Nyl.; Dirinaria confluens (Fr.) Awasthi
{Physcia confluens Mitt.}; Glyphis verruculosa Zahlbr. {Glyphis verrucosa Knight}; Leptogium
cyanescens (Rabenh.) Korb {L. tremelloides L.}; Letrouitia bifera (Ny1.) Hafelln. {Hetero-
thecium biflorum Nyl.}; Ochrolechia pallescens (L.) Massal. {Lecanora pallescens Fr} [?];
Parmentaria ravenelli (Tuck.) Mull. Arg. {Parmentaria havenli Tuck.} [?]; Parmotrema
chinense (Osbeck) Hale & Ahti [Parmelia perlatus L.]; Pertusaria sp.; Phyllopsora parvi-
folia (Pers.) Mull. Arg. {Psora parvifolia Mull. Arg.}; Physcia caesia (Hoffm.) Furnr.;
Physma byrsinum (Ach.) Mull. Arg.; Pseudocyphellaria aurata (Ach.) Vainio {Sticta aurata
Ach.}; Pyrenula nitida (Weigel) Ach. [?]; Pyxine cocoes (Sw.) Nyl.; Ramalina farinacea
(L.) Ach.; Ramalina glaucescens Krempelh. {R. levodea Nyl. var. fastigiata Mull. Arg.}.
Ramalina siliquosa (Huds.) A.L. Sm. {R. scopulorum Ach.} |?]; Ramalina thrausta (Ach.)
Nyl. {R. thrausta (Ach.) Fr.} [?]; Teloschistes flavicans (Sw.) Norm.; Usnea aurantiaco-
atra (Jacq.) Bory {Evernia melaxantha Ach.} |?]; Usnea barbata (L.) Weber ex Wigg. [U.
barbata Ach.}; Usnea florida (1L.) Weber ex Wigg. {U. barbata Ach. var. florida (L.) Fr.};
Usnea intercalaris Krempelh.

Since then only scattered individual reports of lichens occurring in Norfolk Island
have appeared in the literature, such as that of Usnea nexilis Motyka and U. propinqua
Stirton (Motyka, 1938), Ramalina arabum (Dill. ex Ach.) Meyen & Flot. (Riedl, 1976) and
from preliminary reports of the present work (Elix and Streimann, 1985).

PRINCIPAL VEGETATION FORMATIONS

The vegetation of the Norfolk Island complex was divided into five major for-
mations: sub-tropical rainforest, open Avaucarta woodland, pastures and foreshores,
weedy forests dominated by Pszdium and Olea, and Phillip Island (Elix and Streimann,
1985; Ovington, 1984). Brief notes are given below for these, together with the dominant
lichen genera occurring in each formation.

Sub-tropical Rainforest

No doubt the Mt Pitt Reserve, with its remaining areas of sub-tropical rainforest,
is the major habitat for lichens on the island and has the richest lichen flora. Dominant
spermatophytes including Avaucaria heterophylla together with varying amounts of native
hardwoods — Elaeodendron cirtipendulum, Nestegis apetala, Rapanea crassifolia and Baloghia
lucida. ‘The understorey comprises smaller trees and sometimes scattered palms
(Rhopalostylis bauert), tree ferns (Cyathea spp.) and the exotics red guava, Psidium Iittorale

PROG. LINN. SOC. N.S.W., 111 (2), 1989

J. A. ELIX AND H. STREIMANN 107

and Citrus limon. Within this reserve, the eastern and south-eastern slopes of Mt Bates
appear to be the richest site for lichens. Corticolous species abound: the richest sub-
strates are the trunks of Elacodendron and the base of the trunks and branches of Araucaria
heterophylla and Citrus limon. The macrolichen genera Heterodermia, Parmotrema, Ramalina,
Teloschistes and Usnea are prominent in the canopy while Coccocarpia, Pannaria, Physma and
Pseudocyphellaria species occur in the wetter areas at the base of trees. Elaeodendron curti-
pendulum is by far the richest substrate for crustose lichens, including many species of
Graphidaceae, Pyrenula and extensive cover by Clathroporina and Megalospora. One of the
most conspicuous lichens of the forest is the beautiful Pseudocyphellaria aurata, with its
brilliant yellow and emerald green (when wet) thallus, which commonly inhabits the
trunks of Cyathea, Citrus and Araucaria.

Open Araucaria Woodland

Open Araucaria woodlands, which occur at sites such as Anson Bay Reserve,
Selwyn Recreation Reserve and Ball Bay Reserve, constitute the second most important
lichen habitat on the island. The Avaucarza trees throughout the island are characterized
by a prodigious growth of Ramalina arabum and Usnea sp. on the canopy branches in par-
ticular, as well as on lower branches. Dirinaria, Pyxine and Physcia species as well as
various Ramalinae, are very abundant and well developed on the branches of most
Araucaria trees, while Xanthoria and Caloplaca species are limited to Araucarza in coastal
situations.

Pastures and Foreshores

Here old fence posts are colonized by a few corticolous lichens (Usnea, Ramalina,
Lecanora sp.). Remnant Araucarza trees which occur in scattered stands or as individuals
are much better substrates, with more species present and larger populations of lichens.
However their communities are impoverished compared with those on woodland or
forest trees of similar age. Exposed rock surfaces in pasture and along the foreshores are
more interesting substrates. Their communities include representatives of Neofuscelza,
Paraparmelia, Parmotrema and Xanthoparmelia plus a variety of microlichen genera includ-
ing Diploschistes, Lecanora s.1. and Lecidea s.|. Interestingly the limestone outcrops near
Emily Bay provide a suitable habitat for several calciphilic species, including a very
unusual Buellia.

Weedy Forest dominated by Psidium littorale and Olea africans

Areas of Mt Pitt Reserve dominated by thickets of Pszdiwm littorale (guava) are
extremely poor in lichens. Pszdium supports no lichens and its dense canopy prevents
growth on soil or other small trees and shrubs. Lichens in these areas are found virtually
only on scattered remnant Avaucaria or other native trees. Olea africans dominates other
drier areas on the Reserve and, like Pszdium, it is a very poor lichen host. Only scattered
depauperate Usnea, Ramalina and Parmotrema species are found on Olea in such areas.

Phillip Island

The Phillip Island lichen flora is rather depauperate because of the rarity of host
trees. The branches of remnant Lagunaria patersonia are moderately well covered with
lichens, but are relatively poor in taxa, mainly species of Dirinaria, Physcia and Pyxine.
However saxicolous lichens are quite well developed in some areas of Phillip Island (e.g.
upper Long Valley), although again the number of species is not large, with Parmotrema
reticulatum being particularly common. Oddities include VTeloschistes flavicans growing on
soil and rocks, usually a corticolous species, and Xanthoria, Caloplaca and Lecidea growing
on consolidated soil, imitating the soil crust lichens of arid inland Australia.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

108

LICHENS OF NORFOLK ISLAND 1

THE LICHEN FAMILY PARMELIACEAE

Representatives of the genera Flavoparmelia, Neofuscelia, Paraparmelia, Parmelia,

Parmelinopsis, Parmotrema and Xanthoparmelia are detailed and a key to the species is given.
A full description of each species (including chemical constituents) and its distribution
are provided. Interestingly all (16) species have vegetative diaspores, illustrating the ease
of dispersal of such units in reaching such isolated oceanic islands, and hence the
effectiveness of such a mode of reproduction.

Artificial Key to the Parmeliaceae in Norfolk Island

cok Co ES OO ey

pDihtalllhuisqlo nO wins iaectoe ht creed dane v at Mah pa ales cco Coe mea coy fe mea Neofuscelia verrucella
dthallusseneysom yellow ence wes ms. er Ameen eer ai aorm ta ithe aren ea 2
‘Bhallusiyellowsereent ss 42 epee RE a eat SA eds Ce ee ae 3
Mahala; eyes ee PEI RE TERIOR Iieiter Re RDN iar. AUR SERN Aya BAY es ORI eh aap OR de ea 5
sBhvallismvithraspaledloweigsuigialces i: ses aeesni eee ra Xanthoparmelia amplexula
dBinallwsawatheat olackgloweryisuitclalce wry a eatee Oe rains ia eras itr ae yes a ae 4

Thallus with cylindrical isidia, medulla white

(OV COMUCCA COMUNE RRA es tyas of place atarai nau chicane Git pradctcs aes gt Xanthoparmelia australasica
Thallus with pustulate isidia, lower medulla

WEllOW=Orralin oe tener iee sae OM Die nem TS ods sak, Be a Flavoparmelia norfolkensis
ILOIOeS Marans ESS Waain I Bimnian ROC! 4 oy cellos C4 ob Ades bose s Webs eobae 6
ILOlnes loroacl, GReAior tnain Zisman lOPOAG! oo. 5 oo obs os Haddagideda Bou doe eo ale J
divallusgisrdiatessaxicoloush ary ean heen el eee Faraparmelia scotophylla
Thalllus joustullate, wsuallhy CormiColoOUs . 2505506050550 50005 Parmelinopsis spumosa
Ujnoer swriace joseucloovplncllaite 2200 ¢a5 ccs scoues concourse va Farmelia erumpens
Wippemsunacelackimegpscudocypiellacier ere ieee tetera re eae 8
Athallususidiateromsonediatea.2s ues +c oleate hha Maile a Wait meee eee 9
Thallus lacking soredia or isidia, but with flaking upper

COOKE Was Ree eects Iaido tiirsbie nealing sais ade Parmotrema austrocetratum
Pla ANI SHISTGII AWC teens, eset eee ute an stacey cout an UR Ae ivydtryin e (outa Cee gt ee eae ea 10
ablvalltisnsone diate mr tabs sees ie as occa tican eeuaret Ae ee et sek ar ene: eee eae 11
Winaillus culiaice, rmechuilley oe yell, (C= 4.05 co caccsesa0ccue Farmotrema crinitum
Wells eeuliate, waecohuilley Ka (Cre Cl ao eo sca bo ooo soar soc Parmotrema tinctorum
Wippermsunlacemencularelyacrackedinem em aera eae rene Parmotrema reticulatum
Wippemsinracelackine retieulatereracks) sy em) sais einen ee eee te 2.
IMieclWNAIC syalllow Or leas WEllOWReGl os eg oss acco nanesbsaoeacbseessors IS
IVA TCSX I Baus) Se ig ay eas ir Sa ete rea eee Ly Htoe sR NSA Ae el 14
Medulla K+ yellow-red, containing salazinic acid ......... Parmotrema cristiferum
Medulla K+ yellow, containing stictic acid ...........4.... Farmotrema chinense
Medulla C+ rose, containing gyrophoric acid .......... Farmotrema sancti-angelit
Mecha Crease ren). ae ena ey a ee oe a eee eee ie eo ee one 15
Thallus ciliate, medulla P-, UV+, containing alectoronic

GIG EY ahs MoMA ihe Chee te ee shi Lo sad Team bea aes RE Farmotrema rampoddense
Thallus eciliate, medulla P+ brick-red, UV-, containing protocetraric

ACI. n ihe tigls Shs Hid MANET he Manny ini hae Sane CINE Ne ee eR oo Parmotrema gardneri

PROG. LINN. SOG. N.S.W., 111 (2), 1989

J. A. ELIX AND H. SYREIMANN 109

Specimens Examined

Collectors and location of specimens examined are as follows: JAE were collected
by J. A. Elix and H. Streimann and are held in ANUC; HS were collected by H. Stretmann
and are held in CBG with duplicates distributed as indicated.

Collection site details (for map see Fig. 3):
SIE open Araucaria woodland, Picnic Area, end of Martins Road, 29°03’30"S,
167°59’E, 80m, 1.x11.1984.

29°S
NORFOLK ISLAND
4 NEPEAN ISLAND
29°05" 0 1 2km
N
@
PHILIP ISLAND
167°55S° 168°E

Fig. 3. Collection sites on Norfolk Island.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

110

Swe
S3:

S4:

S5s

S6:

SWE

S8:

S9:

S10:

Sil:

S12:

S13:

S14:

S15:

S16:

S17:
S18:

S19:
S20:
$21:
S225
S85
$24:

S25:

LICHENS OF NORFOLK ISLAND 1

pasture, Cemetery road, Kingston, 29°03’30"S, 167°58’E, 15m, 2.xii.1984.
Araucaria heterophylla-dominated grassland, Bloody Bridge, Cemetery Road,
DONS HOGS, UOMO DS 18, Oren, Zs e4e,

mixed exotic and Araucaria woodland, Rocky Point Reserve, 29°03’S,
loo 20) 46 Ome Zax O82

mixed sub-tropical rainforest, near Broken Pine, Mt Pitt Reserve, 220-240m,
DASA RO Sy MOP DO> ANE, Zoek,

stand of Araucaria heterophylla along the margin of grassland and forest, Captain
Cook Memorial, Duncombe Bay, 29°00'S, 167°56’30”E, 100m, 3.x11.1984.
regrowth forest, Just south of the Captain Cook Memorial, Duncombe Bay,
UOX00 20S, NOPD HO 18, UOOran, Src We.

mixed sub-tropical rainforest, Filmy Fern Trail, Mt Pitt Reserve, 29°01'20”S,
OPO 44018, Oro, Joan Oye,

open woodland with kikuyu grass, Mt Pitt Reserve, track at end of Selwyn Pine
IRGC ZOOS, O76" KOE, BilSvan, Sosa es,

Olea africans-dominated valley, Upper Long Valley, Phillip Island, 29°07’30’S,
SIDI Vd., BOiam, Fess".

open Araucaria woodland, Point Blackbourne Reserve, end of Two Chimneys
IRGaiel, 2°03 S, G79 1, Sian, Z.sou NM OSyee.

mixed sub-tropical rainforest, Mt Bates summit trail, Mt Pitt Reserve,
29200 C3 0S 167,296) 3073 00m losxan O84

mixed sub-tropical rainforest, track from Red Road to Mt Bates, Mt Pitt Reserve,
DOYO" AOS, IGT SG 440 1, P20 ma, Oasys.

mixed sub-tropical rainforest, King Fern Valley, Mt Pitt Reserve, 29°01’S,
NOP 2O 18, Aodhan, ovis,

grassland with scattered Araucaria heterophylla, Flat Rock Bay Picnic Area, Anson
Bay IRo@ael, ZO “SOS, IO 5H 18, Oi, Jasin OSH,

Psidium- and Olea-infested lowland forest, track at end of Selwyn Pine Road,
PIONS), MOP SG 308, 200mm, Bsr,

rocky foreshore, Ball Bay Reserve, 29°03’S, 167°59’E, 2m, 8.x1i.1984.

rocky cliffs at north-west point, Duncombe Bay, 29°00’S, 167°55’30”E, 50m,
Oral O34.

grassland with scattered Araucaria heterophylla, near Jacobs Rock, Anson Bay
Re@aG!, LGC", IG 7° D518, S00, Osan ISSHe,

Araucaria-dominated grassland, Bumboras Reserve, 29°03'30"S, 167°56’20”E,
Oren, Osan iSe4e.

open pasture, “(he Cockpit, Cascade Creek Valley, 29°01’S, 167°58°E, 35m,
9) SAMOS,

open woodland, Prince Philip Drive, 29°01’S, 167°58’E, 35m, 9.xi1.1984.

open pasture, Kingston, 29°03’30"S, 167°57'30"E, 8m, 9.x11.1984.

regrowth rainforest, Mt Pitt Reserve, just south of summit of Mt Pitt, 29°01’S,
167°56’E, 230m, 10.x11.1984.

mixed sub-tropical rainforest, Bird Rock ‘Track, Mt Pitt Reserve, 29°00'30"S,
GJ D7 ld, AlOran, Osan Gey,

FLAVOPARMELIA Hale

Flavoparmelia norfolkensis Elix & Streim. sp. nov. (Fig. 4)
Species cum thallo ut in Flavoparmelia euplecta sed ad hac specie isidiis cylindricis, coral-
loideis demum inflatis, apicibus saepe erumpentibus sed esorediosis differt.

Type: Norfolk Island. On Elaeodendron in open woodland in forest clearing, Mt Pitt

PROC. LINN. SOG. N.S.W,, 111 (2), 1989

JA. ELIX AND. STREIMANN 11]

INeschvcminackeatendrot Selwyn PimelRoad 29201 “Ssi167°d6 307 215mm, A. lax
18738 and H. Strermann, 8.xi1.1984 (CBG-holotype; MEL, US-isotypes).

oy ng

we,
pide So,
x

Fig. #4. Holotype of Flavoparmelia norfolkensis.

Thallus foliose, corticolous or saxicolous, adnate, pale yellow-green, to 5cm in
diameter.

Lobes irregular, 1.0-3.0(-5.0)mm wide, becoming laterally imbricate, + contiguous,
plane, rotund at the apices. Upper surface plane to + undulating, dull to shghtly shiny,
with reticulate, white maculae towards the apices, isidiate; isidia laminal, cylindrical at
first and becoming coralloid, ultimately becoming inflated at the apices and erumpent-
pustulate, hollow within (dactylate), very rarely with traces of granular soredia; medulla
white for the most part, but with a yellow-orange zone adjacent to the lower cortex.
Lower surface black with a very narrow, naked, brown rim, sparsely rhizinate, rhizines
short, simple or tufted at the apices, concolorous with the lower surface. Apothecia up
to 5mm in diam., disc strongly concave, dark cinnamon brown, dull; margin persistent,
strongly inrolled, pustulate-isidiate; spores ellipsoid, 19-21 x 8-94m. Pyenidia rare,
black, punctiform, immersed; conidia weakly bifusiform, 4-6 x lum.

Chemistry: Cortex K-; medulla K+ pale brown, C-, P+ brick red; pigmented lower
medulla K+ claret; containing usnic acid, protocetraric acid (major), atranorin (+
trace), euplectin (minor), virensic acid (trace), gyrophoric acid (trace), skyrin (trace).

PROG. LINN. SOG. N.S.W., 111 (2). 1989

112 LICHENS OF NORFOLK ISLAND 1

This species is characterized by the adnate, yellow to yellow-green thallus, the
pustulate isidiate upper surface and a medulla pigmented orange (adjacent to the lower
cortex) with the anthraquinone, euplectin. In many respects this new species resembles
the common Australian species, / euplecta (Stirton) Hale, as both have a similar,
pigmented lower cortex and analogous chemistry. However the upper surface of F
euplecta is sorediate, with laminal, erumpent-pustulate soralia, which are subcapitate at
first but coalesce and spread over the upper surface as granular soredia, whereas F
norfolkensis 1s isidiate with the cylindrical isidia ultimately becoming coralloid, inflating
at the apices and bursting open, but remaining esorediate for the most part. This species
is quite common throughout the island.

Specimens Examined:

NORFOLK ISLAND. On tree trunk and dead log, $5, JAE 18333, 18352; on Elaeodendron,
S7, JAE 18379; on canopy of fallen tree, $8, JAE 18421; on Elaeodendron, S9, JAE 18440,
on Citrus limon, S12, JAE 18580; on fallen Araucaria heterophylla, S12, HS 34354; on dead
treelet branch, S13, HS 34396; on Citrus limon, S14, JAE 18669; on volcanic rocks, S20,
JAE 19269; on volcanic rocks, $21, JAE 18787, on Citrus limon, S24, JAE 18815; on rock
shaded by old building, $23, HS 34806 (US).

PHILLIP ISLAND. On volcanic rocks with a southerly aspect in the open, S10, JAE 16501.

NEOFUSCELIA Esslinger

Neofuscelia verrucella (Essl.) Essl., Mycotaxon 7: 53 (1978).

Parmelza verrucella Essl., in C. F. Culberson, W. L. Culberson and T. L. Esslinger, Bryolo-
ms S02 132 (O77).

Type: Australia. Victoria: Rock Bore, about 33km NNW of Murrayville, Dahl
(O-holotype).

Thallus saxicolous, moderately to loosely adnate, yellow-brown to dark brown, 4-6cm
in diam. Lobes irregular, 1.0-2.5mm wide, often markedly imbricate to entangled.
Upper surface smooth, shining at margins, wrinkled and cracked in the centre, emacu-
late, + lightly pruinose, moderately to densely isidiate; isidia laminal, cylindrical, sim-
ple or branched, to 1.5mm tall; medulla white. Lower surface black, smooth and
shining at margins, matt and wrinkled at centre, sparsely to moderately rhizinate, rhi-
zines simple, black. Apothecia rare, to 2mm in diam., sessile, deeply concave when
young, becoming + flat with age, disc smooth, light to dark-brown, margins entire or
sparsely isidiate; spores broadly ellipsoid, 8-9 x 4-6 um. Pycnidia not seen.

Chemistry: Cortex K-- HNO; + dark blue-green; medulla K-, C-, KC- or KC + faint
rose, P-; containing divaricatic acid (major), nordivaricatic acid (minor).

A common and widespread species in Australia, also occurring in both islands of
New Zealand and South Africa. Rare in Norfolk Island.

Specimen Examined:
NORFOLK ISLAND. On volcanic rocks, S21, JAE 18791.

PARAPARMELIA Elix & Johnston

Paraparmelia scotophylla (Kurok.) Elix & Johnston, Mycotaxon 27: 281 (1986).

Farmelia scotophylla Kurok., in Kurokawa and Filson, Bull. natn. Sci. Mus. Tokyo B, 1: 46
(1975).

Type: On rocks, Ardglen Gap, Liverpool Range, 8.5km north of Murrurundi, Great
Dividing Range, New South Wales, S. Kurokawa 5174, 28.x.1965 (TNS-holotype;
MEL-isotype).

PROG. LINN. SOG. N.S.W,, 111 (2), 1989

J. A. ELIX AND H. STREIMANN 113

Thallus saxicolous, adnate, mineral grey but blackening with age, 5-10 (-20)cm diam.
Lobes irregular to sublinear, 1.0-3.0mm wide, slightly imbricate, apices subrotund.
Upper surface smooth, shining at margins, flat to slightly convex, becoming cracked in
the centre, emaculate, moderately to sparsely isidiate; isidia laminal, cylindrical,
simple or sparingly branched, to 0.2mm tall; medulla white. Lower surface pale to light
brown, sparsely to moderately rhizinate, rhizines simple, concolorous. Apothecia rare,
to 6mm in diam., substipitate, deeply concave, disc smooth, pale brown, margins thin,
involute, isidiate; spores 7-11 x 5-8 ym. Pycnidia not seen.

Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange;
containing atranorin, salazinic acid (major) and consalazinic acid.

A common and widespread species in Australia, also occurring in both islands of
New Zealand. Rare on Norfolk Island.

Specimen Examined:
NORFOLK ISLAND. On volcanic rocks, $21, JAE 18793.

PARMELIA Acharius

Parmelia erumpens Kurok., Lich. rar. Critic. Exsicc. no. 74 (1969)

[Based on Parmelia tenuirama J. D. Hook. & Taylor f. corallina Mull. Arg.]

Parmelia tenuirima J. D. Hook. & Taylor f. corallina Mull. Arg., Flora, Jena, 66: 46 (1883).
Type: Gippsland, Australia, Starling (G-lectotype; UPS, US-isolectotypes).

Thallus corticolous or saxicolous, adnate to loosely attached, pale greenish to light
mineral-grey, 8-20cm in diam. Lobes short, subirregular to apically rotund, imbricate,
2-8mm wide. Upper surface shiny, plane, white-reticulate at first but becoming con-
spicuously cracked to the margin, pseudocyphellae effigurate, 0.2-1.0mm long, some-
what raised, dense, fusing into a reticulate network over the whole surface, sorediate;
the soredia coarse and isidioid, often bursting apically, forming dense marginal and
laminal soralia and/or extended, granular, coralloid-isidioid outgrowths; medulla white.
Lower surface black, moderately rhizinate, rhizines simple or squarrosely branched at
maturity, 1-2mm long, black. Apothecia rare, stipitate, to 15mm in diameter, disc dark
brown, concave at first then flattening, margin inrolled at first but splitting radially at
maturity, the amphithecium reticulately cracked, pseudocyphellate, sorediate; spores
ellipsoid, 10-12 x 6-8um. Pycnidia scattered, punctiform, black, conidia cylindrical to
weakly bifusiform, straight, 5-7 x lum.

Chemistry: Cortex K+ yellow; medulla K+ yellow becoming deep red, C-, P+ red-
orange; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid
(minor), lobaric acid (minor), protocetraric acid (+ trace).

P. erumpens is a widespread species on rocks and trees in coastal and hinterland
areas of Australia, both islands of New Zealand and Lord Howe Island. It also occurs in
South Africa, India, Indonesia, Taiwan and Japan. This species is scattered on both
rocks and trees in Norfolk Island and is easily recognized by the deeply reticulate-
fissured cortex, the abundant coarse, pustular soredia and the lack of cilia. Uncommon
in Norfolk Island.

Specimens Examined:
NORFOLK ISLAND. On exposed volcanic rocks, $2, JAE 16316, HS 31746; on Elaeoden-
dron, 55, JAE 18310; on tree, S7, JAE 18360, 18385; on Elaeodendron, S16, JAE 18737.

PARMELINOPSIS Elix & Hale
Parmelinopsis spumosa (Asahina) Elix & Hale, Mycotaxon 29: 243 (1987).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

114 LICHENS OF NORFOLK ISLAND 1

Farmelina spumosa (Asahina) Hale, Phytologia 28: 483 (1974).

Farmelia spumosa Asahina, J. Jap. Bot. 26: 259 (1951).

Type: Higashi-Murayama, Kita-Iama-gun, Prov. Masashi, Japan, Asahina (1NS-
lectotype).

Thallus corticolous, adnate, fragile, pale mineral grey to pale olive grey, 2-6cm in diam.
Lobes sublinear, narrow, 0.5-2.0mm wide, ciliate, the marginal cilia distinct and evenly
dispersed, ca. 0.5mm long. Upper surface plane, emaculate, continuous, densely
pustulate-isidiate; pustules laminal, bursting open but sparingly or not sorediate;
medulla white or faintly yellow. Lower surface smooth, shiny, black, moderately
rhizinate, rhizines simple or furcate, short, ca. 0.1-0.2mm long. Apothecia very rare,
laminal, adnate, concave, 1-3mm in diam., disc brown, imperforate, margin and
amphithecium pustulate; spores ellipsoid, 12-14 x 7-8um. Pyenidia not seen.

Chemistry: Cortex K+ yellow; medulla K-, C+ rose, KC + red, P-; containing atra-
norin, chloroatranorin, gyrophoric acid (major), 5-O-methylhiascic acid (minor),
umbilicaric acid (minor).

P. spumosa 1s a cosmopolitan species which is widespread throughout the tropical
and sub-tropical areas of the world, but is much less common at temperate latitudes. In
Australasia it is common in Australia, Papua New Guinea and New Zealand. Exposed
trees at the forest margins or isolated individuals are the preferred habitat. Rare in

Norfolk Island.

Specimens Examined:
NORFOLK ISLAND. On Citrus limon, S12, JAE 16584; on Elaeodendron, 322, JAE 18798; on
Eiaeodendron, S25, JAE 18639, 10.x11.1984.

PARMOTREMA Hale

Parmotrema austrocetratum Elix & Johnston, Mycotaxon 31: 495 (1988)
Type: New Zealand. North Island. On tree trunk in remnant forest, Burgess Park, New
Plymouth, 60m, /. A. Elix 4645, 7v.1980 (CHR-holotype, CBG-1sotype).

Thallus corticolous or saxicolous, loosely attached, light mineral-grey, 6-12cm in diam.
Lobes rotund, imbricate or not, 10-20(-30)mm wide, apices often laciniate, the lacinae
flat or convex, 0.5-1.5mm wide, 1-5(-8)mm long, lobes moderately ciliate, the cilia 0.2-
1.0(-1.5)mm long, simple or sparingly branched. Upper surface white-reticulate at first
but becoming conspicuously cracked to the margin, developing raised closed dactyls,
dactyls + curved, laminal, ultimately becoming fused and forming laminal ridges, the
older parts of the thallus eventually becoming cracked-areolate, with the areolae flaking
off and exposing the white medulla in eroded areas; lacking soredia and isidia. Lower
surface black with a bare, brown marginal zone 1-2mm wide, moderately to densely
rhizinate, rhizines simple or sparsely branched, slender, black. Apothecia rare,
stipitate, to 10mm in diameter, disc perforate, pale tan, concave at first then concave-
distorted, margin eciliate, thin, + stellate-cracked; spores ellipsoid, 12-16 x 8-9um.
Pycnidia scattered, punctiform, black, conidia filiform, 9-16 x 1m.

P. austrocetratum is a widespread species on rocks and trees in coastal and hinterland
areas along the sub-tropical and tropical east coast of Australia and the north island of
New Zealand. Uncommon in Norfolk Island.

PROG. LINN. SOC. N.S.W., 111 (2), 1989

J. A. ELIX AND H. SUREIMANN 115

Specimens Examined:
NORFOLK ISLAND. On Cyathea stem, 55, HS 31905; on Elaeodendron, 88, JAE 18439, HS
32190 (B, H, US); on Elaeodendron, S16, JAE 18742.

Parmotrema chinense (Osbeck) Hale & Ahti, Taxon 35: 133 (1986).

Lichen chinensis Osbeck, Dagb. Ostindisk resa,: 221 (1757).

Type: Specimen and pl. 20, fig. 39B, Dillenius, Host. Musc.: 147 (1742) (OXF
lectotypotype).

Lichen perlatus Huds., Fl. angl.: 448 (1762), (incl. type of L. chinensis Osbeck).

Farmelia perlata (Huds.) Ach., Meth. Lich.: 216 (1803).

Parmotrema perlatum (Huds.) Choisy, Bull. mens. Soc. linn. Lyon 21: 174 (1952).

Thallus corticolous, moderately to loosely adnate, membranaceous to coriaceous, pale
mineral grey to whitish-grey, 4-15cm in diam. Lobes irregular, 3-8mm wide, in part
rounded and deeply crenate, in part irregularly incised and laciniate, imbricate,
sparingly to moderately ciliate, cilia 0.2-3.0mm long, + branched. Upper surface dull,
smooth, sorediate; soralia submarginal, causing lobe margin to become revolute and
suberect, ultimately appearing labriform, soredia granular; medulla white. Lower sur-
face black, shining, with a broad, brown, naked marginal zone, moderately rhizinate,
rhizines simple, to 2mm long. Apothecia very rare, laminal, substipitate, concave, to
7mm in diameter, disc pale brown to cinnamon brown, imperforate, margin thick,
inrolled, sorediate; spores ellipsoid, 25-27 x 16-18um. Pyenidia not seen.

Chemistry: Cortex K+ yellow; medulla K+ yellow, C-, P+ orange-red; containing
atranorin, chloroatranorin, stictic acid (major), constictic acid (major), cryptostictic
acid (trace), menegazziaic acid (trace), norstictic acid (trace).

P. chinense 1s a cosmopolitan species which is widespread throughout the tropics and
temperate areas. In Australasia it is common in Australia and New Zealand, but also
occurs in Fiji and Papua New Guinea. It is very rare on Norfolk Island, and although it
was the first ‘“Farmelia’ reported for the island (Endlicher, 1833), this may well have
referred to P. reticulatum, a morphologically similar and common species.

Specimen Examined:
NORFOLK ISLAND. On treelet stem, S24, HS 34830 (US).

Parmotrema crinitum (Ach.) Choisy, Bull. mens. Soc. linn. Lyon 21: 175 (1952).
Parmelia crinita Ach., Syn. Lich.: 196 (1814).
Type: North America, Muhlenberg (H-holotype).

Thallus corticolous or saxicolous, coriaceous, adnate, pale mineral grey to grey-green,
5-6cm in diameter. Lobes subirregular, crenate or irregularly incised, imbricate, 2-
5mm wide, the margin ciliate, cilia simple or branched, 0.5-3.0mm long. Upper surface
plane, emaculate, smooth to rugose with age, cortex fragile, isidiate; isidia laminal and
marginal, short-cylindrical at first, + becoming coralloid, granulose or occasionally
dissolving into soredia, often ciliate at the apices; medulla white. Lower surface black,
with a narrow brown, naked marginal zone, densely rhizinate, rhizines slender, simple,
to 1.0mm long. Apothecia and pycnidia not seen.

Chemistry: Cortex K+ yellow; medulla K+ yellow, C-, P+ orange; containing atra-
norin, chloroatranorin, stictic acid (major), constictic acid (minor), cryptostictic acid
(trace), norstictic acid (trace), menegazziaic acid (+ trace), connorstictic acid (+
trace).

P. crinitum is a cosmopolitan species, widespread in humid habitats in temperate
and tropical regions. In Australasia this species 1s common in Australia, New Zealand
and Papua New Guinea. Common in Norfolk Island.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

116 LICHENS OF NORFOLKISLAND 1

Specimens Examined:

NORFOLK ISLAND. On base of Araucaria heterophylla, 51, JAE 18133; on base of Araucaria
heterophylla, S4, JAE 16216, 16246, 16323, HS 31821 (US); on tree and dead branches,
S5, JAE 18276, 16293; on Araucaria stem, S6, HS 32018 (US); on Elaeodendron S7, JAE
18584; on Lagunaria, $8, HS 32181 pr. p.; on Campsis grandiflora, S9, JAE 18469; on base of
Araucaria heterophylla, Sil, JAE 18513; on Citrus limon, S12, JAE 18579, 18581, 18588,
18590; on treelet stem, S12, HS 34376; on palm and Citrus limon, S14, JAE 18670, 18679,
18692, HS 34515; on volcanic rocks, $17, JAE 16749; on volcanic rocks, $20, JAE 19268;
on Citrus limon, S24, JAE 18812, 18817.

Parmotrema cristiferum (Taylor) Hale, Phytologia 28: 335 (1974).
Farmelia cristifera Yaylor, Hooker’s Lond. J. Bot. 6: 165 (1847).
Type: Calcutta, Wallich (FH-lectotype; BM-isolectotypes).

Thallus corticolous, adnate to loosely adnate, coriaceous, pale mineral grey, 3-10cm in
diam. Lobes irregular, 5-20mm wide, rotund at the apices, entire or weakly crenate,
imbricate or subascending at the margins, main lobes eciliate, lateral lobes and lobe
axils not or sparingly ciliate, cilia 0.5-1.5mm long. Upper surface dull, emaculate, con-
tinuous, sorediate; soralia mainly marginal on lateral lobes, sorediate lobes more or less
ascending, soredia granular; medulla white. Lower surface smooth, shiny, black, with a
broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, short, ca 0.1-
0.2mm long, coarse. Apothecia very rare, laminal, adnate, concave, to 3mm in
diameter, disc brown, imperforate, margin thick; spores ellipsoid, 25-30 x 13-15ym.
Pycnidia not seen.

Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange-red;
containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (+
minor).

P. cristiferum is a cosmopolitan species that is widespread throughout tropical and
sub-tropical areas. In Australasia and Oceania it is common in Australia, Fiji, Papua
New Guinea, Pitcairn Island, Raratonga and New Zealand. Common in Norfolk
Island.

Specimens Examined:

NORFOLK ISLAND. On base of Avaucaria heterophylla, S1, JAE 18138; on dead branches
and shrubs, $5, JAE 18279, 18282, 18301, HS 31908; on Elaeodendron stem, S7, HS
532061; on Elaeodendron, S8, JAE 18420, 18427; on Elaeodendron, S9, JAE 18444; on Citrus
lumon, S12, JAE 18581, 18588, 18590, HS 34264 (B, H, US); on dead treelet stem, S13,
HS 34397, 34414 (US); on palm, S14, JAE 18692; on treelet stem and stump, S14, HS
34515, 34516, on crown of Araucaria, S15, HS 34596; on Elaeodendron, S16, JAE 18735
(Lich. Australasict Exsicc. Fasc. 4: 88); on palm stem and dead tree, S16, HS 34633, 34674,
34675 (US); on Citrus limon, S24, JAE 18812.

Parmotrema gardneri (Dodge) Sérusiaux, Bryologist 87: 5 (1984).
Farmelia gardneri Dodge, Ann. Mo. bot. Gdn. 46: 179 (1959).
Type: Brazil, Gardner (F H-holotype).

Thallus corticolous, adnate to loosely adnate, coriaceous, pale mineral grey, 4-6cm in
diam. Lobes irregular, 8-15mm wide, rotund at the apices, imbricate or subascending at
the margins, crenate, irregularly incised or sublaciniate, eciliate or rarely sparingly
ciliate, cilia 0.2-0.5mm long. Upper surface dull, emaculate, becoming rugose with age,
continuous or + cracked, sorediate: soralia marginal, linear along the margins to sub-
capitate on somewhat revolute lobes or short marginal laciniae, sometimes spreading
submarginally, soredia granular; medulla white. Lower surface wrinkled, black, with a

PROG. LINN. SOG. N.S.W., 111 (2), 1989

J. A. ELLX AND H. STREIMANN 117

broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, short, ca. 0.1-
0.2mm long, slender. Apothecia rare, laminal, adnate, concave, to 3mm in diameter,
dise dark brown, imperforate, margin thick, eciliate, margin and amphithecium soredi-
ate; sores ellipsoid, 18-22 x 8-10um. Pyenidia rare, punctiform, conidia sublageniform,
6-7 x lum.

Chemistry: Cortex K+ yellow; medulla K+ pale brown, C-, P+ brick-red; containing
atranorin, chloroatranorin, protocetraric acid (major), unknown fatty acids (+ minor).

The presence of the marginal soralia and medullary protocetraric acid plus the
absence of cilia distinguish this from other species of Farmotrema on the island. It is a pan-
tropical species known from Africa, Australia, Papua New Guinea and South America.
Rare in Norfolk Island.

Specimen Examined:
NORFOLK ISLAND. On Araucara trunk, S1, JAE 18156.

Parmotrema rampoddense (Ny1.) Hale, Phytologia 28: 338 (1974).
Parmelia rampoddensis Ny1., Acta Soc. Sci. Fenn. 26: 7 (1900).
Type: Ramboda, Ceylon, Almquist (H-NY L 35555-holotype; S-isotype).

Thallus corticolous, loosely adnate, coriaceous, pale grey to mineral grey, 10-20cm in
diam. Lobes irregular, 5-20mm wide, rotund at the apices, crenate, ciliate, cilia con-
spicuous, 3-6mm long, simple or bifurcate. Upper surface dull, emaculate, rugulose
towards the centre, sorediate; soralia mainly marginal, linear, sometimes spreading
submarginally, soredia farinose; medulla white, often pigmented orange-red adjacent to
the lower cortex. Lower surface smooth, shiny, black, with a broad, brown, naked
marginal zone, sparsely rhizinate, rhizines simple, to 6.0mm long, slender. Apothecia
not seen in Norfolk Island material. Hale (1965) reports apothecia 3-10mm in diam.,
disc imperforate, amphithecium sorediate and spores 10-12 x 6-7um. Pyenidia not seen.

Chemistry: Cortex K+ yellow; medulla K-, C-, KC + red, P-; pigmented lower medulla
K+ purple; containing atranorin, chloroatranorin, alectoronic acid (major), a
collatolic acid (major or minor), skyrin (+).

P. rampoddense is a common and widespread pantropical species. In Australasia it is
known from Australia and Papua New Guinea. Rare in Norfolk Island.

Specimens Examined:
NORFOLK ISLAND. On fallen branch; $5,/AE 16/25; on treelet stem, S14, HS 34532.

Parmotrema reticulatum (Taylor) Choisy, Bull. mens. Soc. linn. Lyon 21: 175 (1952).
Farmelia reticulata Vaylor, in Mackay, Fl. Hibern. 2: 148 (1836).
Type: Ireland, County Kerry, near Dunkerron (Ff H-holotype).

Thallus corticolous or saxicolous, loosely adnate, membranaceous to coriaceous, pale
mineral grey to grey-green, 4-20cm in diam. Lobes irregular, 5-15mm wide, in part
rounded and deeply crenate, in part irregularly incised and laciniate, imbricate or
subascending at the margins, sparingly to moderately ciliate, cilia 0.2-3.0mm long.
Upper surface dull, reticulately maculate and cracked, sorediate; soralia marginal,
linear along the margins to subcapitate, commonly on short marginal laciniae, some-
times submarginal and punctitorm, soredia granular; medulla white. Lower surface
black, rhizinate or papillate to the margins or with a brown, naked marginal zone,
densely rhizinate, rhizines simple or squarrose, to 2mm long, slender. Apothecia rare,
submarginal to laminal, substipitate, concave, to 8mm in diameter, disc pale to mid-
brown, imperforate or narrowly perforate, margin thick, eciliate, margin and amphi-
thecium sorediate; spores ellipsoid, 13-18 x 8-11m. Pyenidia rare, punctiform, conidia
filiform, 12-16 x 1.0-1.5um.

PROC. LINN. SOG. N.S.W., 111 (2), 1989

118 LICHENS OF NORFOLK ISLAND 1

Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange-red;
containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (+
minor).

P. reticulatum 1s a very common and highly variable species. Specimens growing in
drier, exposed sites tend to become coriaceous, have weakly developed maculae and
often submarginal soralia, while those from moist, shady habitats are usually membran-
aceous, have marginal soralia and well-developed maculae. P. retzculatum is a cosmopoli-
tan species, widespread throughout the tropics and temperate areas. In Australasia it is
common in Australia, New Zealand and Papua New Guinea. Very common in Norfolk
Island and occasional in Phillip Island.

Specimens Examined:

NORFOLK ISLAND. On Araucaria trunk, S1, HS 31716, 31723, 31728, 51734 (US); on
exposed boulder, S2, HS 31748 (H, US); on small rock outcrop, $3, HS 51768, 31769 (B,
H, US); on Grevillea robusta, and Melia, $4, JAE 18222, 16231, 18250, HS 31791, 31839
(US); on dead branch, $5, JAE 18308; on rocks, $6, JAE 18371, 18374, HS 31997, 32022
(US); on old wooden steps, $8, HS 32169 (H, US); on Cyathea stem, S14, JAE 16665, HS
34557 (US); on Araucaria trunk, S15, HS 34578A, 345868 (US); on volcanic rocks, S17,
JAE 18745, 18746, 18837, HS 34695, 34699, 34707 (US); on volcanic rocks, S18, JAE
18764, HS 34736; on Lagunana patersonia trunk, S19, JAE 187768; on volcanic rocks, S20,
JAE 16833 (Lich. Australasics Exsicc. Fasc. 4: 89), 186834, 19267, HS 34769 (1, US); on
shaded rocks, S21, HS 34782.

PHILLIP ISLAND. On south-facing volcanic rocks in the open, S10, JAE 18502, 18503,
AS 32204.

Parmotrema sancti-angelu (Lynge) Hale, Phytologia 28: 339 (1974).

Parmelia sancti-angelu Lynge, Ark. Bot. 13: 35 (1914).

Type: Santo Angelo, Rio Grande do Sul, Brazil, 25 January 1893, G. A. Malme
(S-lectotype).

Thallus corticolous, adnate to loosely adnate, membranaceous to coriaceous, pale grey
to pale grey-green, 3-10cm in diam. Lobes irregular, 5-15 mm wide, rotund at the
apices, crenate, often deeply divided and with ascending margins, ciliate, cilia slender,
1.0-3.5mm long, simple or bifurcate. Upper surface dull, emaculate, wrinkled or rugose
towards the centre, sorediate; soralia mainly marginal, linear, sometimes spreading
submarginally, soredia farinose; medulla white. Lower surface smooth, shiny, black,
with a broad, brown or mottled, naked marginal zone, densely rhizinate, rhizines
simple, elongate, to 2mm long, slender. Apothecia not seen in Norfolk Island material.
Hale (1965) reports apothecia imperforate, spores 13-18 x 7-10um. Pyenidia rare,
conidia weakly sublageniform, 6-8 x lum.

Chemistry: Cortex K+ yellow; medulla K-, C+ pale red, P-; containing atranorin,
chloroatranorin, gyrophoric acid (major), lecanoric acid (+ trace).

P. sancti-angelit is acommon and widespread pantropical species. In Australasia it is
known from Australia and Papua New Guinea. Rare in Norfolk Island.

Specimen Examined:

NORFOLK ISLAND. On Elaeodendron, 55, JAE 18349.

Parmotrema tinctorum (Despr. ex Nyl.) Hale, Phytologia 28: 339 (1974).
Farmelza tinctoria Despr. ex Nyl., Flora, Jena 55: 547 (1872).
Type: Canary Islands, Despréeaux (H-NYL 35365-holotype).

Thallus corticolous or saxicolous, loosely adnate, membranaceous to coriaceous, pale
grey to grey-green, 10-30cm in diam. Lobes irregular, 10-20mm wide, rotund at the

PROG. LINN. SOG. N.SW., 111 (2), 1989

J. A. ELIX AND H. STREIMANN 119

apices, entire or crenate, eciliate. Upper surface shiny, becoming dull towards the
centre, emaculate, cortex sometimes cracking and flaking, isidiate; isidia sparse to
abundant, laminal and eventually marginal, confluent or in scattered groups, brown
tipped or concolorous with the thallus, simple or branched, thin and cylindrical or
coarse and irregularly inflated, rarely + interspersed with lobules; medulla white.
Lower surface smooth, shiny, black, with a broad, brown, naked marginal zone,
sparsely rhizinate, rhizines simple, short, ca 0.5-2.0mm long, coarse. Apothecia very
rare, laminal, substipitate, concave or + radially split, to 20mm in diameter, disc dark
brown, imperforate or with a small perforation, margin thick, margin and
amphithecium isidiate; spores ellipsoid, 13-15 x 7-8um. Pyenidia rare, conidia filiform,
lGexle Ole opin:

Chemistry: Cortex K+ yellow; medulla K-, C+ red, P-; containing atranorin,
chloroatranorin, lecanoric acid (major), orsellinic acid (trace).

Farmotrema tinctorum is a cosmopolitan species that is widespread throughout
tropical and temperate regions. In Australasia and Oceania it 1s common in Australia,
Fiji, Papua New Guinea, Vanuatu, Samoa and the North Island of New Zealand. Very
common in Norfolk Island and Phillip Island.

Specimens Examined:

NORFOLK ISLAND. On base of Araucaria heterophylla, S1, JAE 18137, HS 31739 (H, US);
on Grevillea robusta, S4, JAE 16221; on Elaeodendron and dead branches, $5, JAE 18285,
18307, 18350; on Araucania trunk, S6, HS 32017, 32055 (US); on canopy of fallen tree,
S8, JAE 16411; on Elaeodendron, 88, HS 32164, 32196 (H, US); on Elaeodendron and Citrus
limon, S12, JAE 16556, 18570; on Araucana trunk, S15, HS 34569 (B, MICH, US); on
Elaeodendron, S16, JAE 18736 (Lich. Australasici Exsicc. Fasc. 4: 90); on rock outcrop, $20,
HS 34660 (US); on tree trunk, $22, HS 36462 (B, US); on Citrus limon, 324, JAE 18616;
on tree in open area, Mt Pitt Reserve, R. Goldsack 4, 25.x11.1981 (ANUC).

PHILLIP ISLAND. On south-facing volcanic rocks in the open, S10, JAE 18499, HS 32225
(B, H, US); on Lagunaria stem, S10, HS 32213, 32225.

VANTHOPARMELIA Hale
Xanthoparmelia amplexula (Stirton) Elix & Johnston, Bull. Br. Mus. nat. Hist. (Bot.) 15:
192 (1986).
Farmelia amplexula Stirton, Trans Proc. Roy. Soc. Victoria 17: 69 (1881).
Type: Australia. Near Brisbane, Queensland, Bailey 262 (BM-holotype).

Parmelia violascens Stirton, Trans Proc. New Zeal. Inst. 32: 77 (1899).
Type: Australia. Grampian Mountains, Victoria, Sullivan (BM-holotype).

Thallus foliose, adnate to moderately adnate on rocks, yellow-green or commonly
blackening towards the centre as a result of the dark-tipped, dense isidia; subirregularly
lobate, 5-10cm in diam. Lobes sublinear-elongate, sparingly imbricate or not so, 0.8-2.0
(3.0)mm wide. Upper surface opaque, emaculate, with sparse to numerous isidia, isidia
cylindrical, simple or branched and coralloid up to 2mm high; medulla white. Lower
surface pale brown to dark brown, sparsely to moderately rhizinate, the rhizines con-
colorous with the lower surface, simple, slender. Apothecia rare, 1.5-6.0mm in diam.;
disc strongly concave to + flat at maturity, dark brown, shining; margin thin, persis-
tent, involute at first, isidiate; spores 7-12 x 5-6um. Pyenidia not seen.

Chemistry: Cortex K-; medulla K-, C-, KC + rose, P-; containing usnic acid, loxodin
(minor), norloboridone (major), unknown (trace).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

120 LICHENS OF NORFOLK ISLAND 1

A common and widespread species in Australia (occurring in all States and Terri-
tories) and both islands of New Zealand. It also occurs in Lord Howe Island and South
Africa. Rare in Norfolk Island.

Specimen Examined:
NORFOLK ISLAND. On treelet stem, $25, HS 349/64.

Xanthoparmelia australasica D. Gall., N.Z. J. Bot. 18: 531 (1980).

Parmelva australasica (D. Gall.) Filson, Aust. J. Bot. 30: 519 (1982).

Type: New Zealand, North Auckland, Karekare Beach, on andesitic conglomerate, 8
October 1977, J. Bartlett (CHR 314047 — holotype; CHR — isotypes).

Thallus foliose, moderately to loosely adnate, yellow-green, 5-12 (-20)cm in diam.
Lobes irregular, 2.0-3.5 (-5.0)mm wide, often markedly imbricate, secondary lobes
similar to the marginal lobes, sometimes building the thallus up into a thick mat, apices
subrotund. Upper surface smooth, shining at margins, wrinkled and cracked in the
centre, emaculate, moderately to densely isidiate; isidia laminal, often forming a dense
areolate crust, robust, broad, cylindrical, extensively coralloid-branched, to 2.5mm tall;
medulla white. Lower surface black with a narrow brown, naked, marginal zone,
smooth and shining at margins, matt and wrinkled at centre, sparsely rhizinate, rhizines
simple, black. Apothecia rare, 2-6 (-8)mm in diam., sessile, deeply concave when
young, becoming shallowly convex with age, disc smooth, red-brown, margins con-
spicuously isidiate, exciple wrinkled, shining, becoming isidiate with age; spores 8-11 x
4-6um. Pycnidia not seen.

Chemistry: Cortex K-; medulla K+ yellow then dark red, C-, KC + red, P+ orange;
containing usnic acid, salazinic acid (major) and consalazinic acid, protocetraric acid
(+ trace), norstictic acid (+ trace).

A common and widespread species in Australia, which occurs in all States and
Territories. It also is found in both islands of New Zealand, the continental United
States and South Africa. Relatively uncommon in Norfolk Island.

Specimens Examined:

NORFOLK ISLAND. On rock in shaded track cutting, S13, HS 543714; on volcanic rocks,
S18, JAE 18765; on volcanic rocks, S21, JAE 18788, 18789, 18790, 18794, 18797, HS
34792 (US).

ACKNOWLEDGEMENTS

We thank the Australian National Parks and Wildlife Service for generous financial
support for our field work on Norfolk Island, Dr G. A. Jenkins who determined the
chemistry of many specimens and Ms Jen Johnston for many constructive suggestions to
this manuscript.

References
Coyne, P. D., 1982. — Day of the rabbits. Studics on plant regeneration on Phillip Island. Geo 4: 30-39.
ELIX, J. A., 1985. — Lichenes Australasici Exsiccatt. Fasc. 4. Canberra: Australian National University.
, and STREIMANN, H., 1985. — Lichens and bryophytes of Norfolk Island. (Unpublished report to the
Australian National Parks and Wildlife Service).
ENDLIGHER, S., 1833. — Prodromus Florae Norfolkicae, sive Catalogus Stirpium quac in Insula Norfolk Annis 1804 et
1805 a Ferdinando Bauer collectae et depictae. Vienna: Fricdrich Beck.
Har, M. E., 1965. — A monograph of Parmelia subgenus Amphigymnia. Contr. U.S. Natn. Herb. 36: 193-358.
Hrwarb, R., 1842. — Biographical sketch of the late Allan Cunningham, Esq. Hooker’ Lond. J. Bot. 1: 107.
Hoart, M., 1965. — Rambler’s Guide to Norfolk Island. Sydney: Pacific Publications.
Jones, J. G., and McDouGALL, I., 1973. — Geological history of Norfolk and Phillip Islands, southwest
Pacific Ocean. J. Geol. Soc. Aust. 20 (3), 239-257.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

J. A. ELIX AND H. STUREIMANN 121

MAIDEN, J. H., 1904. — Vhe flora of Norfolk Island. Proc. Linn. Soc. N.S.W. 28: 692-785.

Moryka, J., 1936-38. — Lichenum gencris Usnea studium monographicum. I. Pars systematica. Leopoli (1.c. Lublin,
Poland).

OVINGTON, J. D., (director), 1984. — Plan of Management Norfolk Island National Park. Canberra: Government
Printer, for Australian National Parks and Wildlife Service.

RiEDL, H., 1976. — Beobachtungen an Ramalina arabum (Ach.) Meyen ct Flot. und Ramalinopsis mannii
(luck.) Follm. ct Hun. Sydowra 28: 134-142.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

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Conicochernes doyleae, anew Australian Species of
the Chernetidae (Pseudoscorpionida: Arachnida)

CLARICE M. A. KENNEDY

KENNEDY, C. M. A. Conicochernes doyleae, a new Australian species of the Chernetidac

(Pseudoscorpionida: Arachnida). Proc. Linn. Soc. N.S.W. 111 (2), 1989: 123-129.

A new species of the genus Conicochernes Beier, C. doyleae, is described from the
Sydney region. It is corticolous, being associated in the sampled area specifically with
the ‘Brush Box, Lophostemon confertus (R.Br.) Peter G. Wilson ct J. T. Waterhouse.
Clarice M. A. Kennedy, School of Biological Sciences, Macquarie University, North Ryde, Australia
2109, manuscript received 16 August 1986, accepted for publication 15 February 1989.

INTRODUCTION

The Chernetidae are the largest family of pseudoscorpions currently including
nearly 100 genera and over 550 species distributed over much of the world (Harvey,
1985). Of these only 12 genera, represented by 26 species, occur in the Australian
region.

The species described herein is a representative of the genus Conicochernes Beier
which is endemic to Australia. Four species are currently known namely: C. brevispinosus
(L. Koch), C. crassus Beier, C. globosus Beier and C. incrassatus (Beier).

Measurements are based on the examination of 10 specimens of each stage made in
accordance with those advocated by Chamberlin (1931). Those in parentheses are the
female and follow those of the male.

Abbreviations for chelal trichobothria and setal formulae follow those devised by
Chamberlin (1931).

Genitalic terminology follows Legg (1974a, 1974b). All specimens are preserved in
spirit.

SYSTEMATIC DESCRIPTION
CHERNETIDAE Chamberlin 1931
Conicochernes Beier 1948
Type species: Chelifer brevispinosus L. Koch 1885.
For synonymy list refer to Harvey 1981: 246.

KEY TOSPECIES OF CONIGCOCHERNES

i orciistalibladcrottlacellumascrrates 26 34.52 ceed Ae Sele ss 2
Shekalistalgolac erotilacellmmypnotsenicatels aay eee) aes Cera 3
2, ENCICASE Z Galles SAAS, GS SOUS ORES =o 3 a se eee see geo C. doyleae sp. nov.
icaleasctaMeyesspotsialseiter wei ay. erry a aeae sn tae: C. incrassatus (Beier)
3, omalahigtaliolacks ot ikereelikunes oyvarneitis 3. 455%s0ebe ues seen sssgeoneesssce-s 4
ANI ollevoles Ont ilavereliiluran jOUMIMAS . oes soc soe aasceaepsavccecae C. globosus Beier
4, Serrulla Sxiemor wile WOl7 lernellae 2. ese cesses: C. brevispinosus (LL. Koch)
SEMAN with Zi Warne 2 so cache no see uscecenseoeass C. crassus Beier

Conicochernes doyleae sp.nov.

(Figs 1-4)
Etymology: This species is named after Suzanne Doyle, a colleague who has been of
great assistance to me in the field.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

124 NEW AUSTRALIAN PSEUDOSCORPION

Fig. 1. Conicochernes doyleae sp.nov., scanning electron micrographs of paratypes: A, right chela, lateral aspect,
male; B, left chela, lateral aspect, female; C, dorsal seta of tergite 4; D, serrula exterior of chelicerac, female;
Da flagellum of left chelicera; E, movable finger of left chelicera.

Holotype: 9, Australian Museum, Sydney (KS 17445).

Type locality: Stanley Street, Chatswood, 10km NW of Sydney G.P.O., N.S.W., lat.
33°48°10"S, long. 151°11’30”E, under bark of Lophostemon confertus (R.Br.) Peter G.
Wilson et J. T. Waterhouse, 17. iv. 1985, C. Kennedy.

Paratypes: same data as holotype, Australian Museum, 40 (KS 17446), 49 (KS
19207), 4 nymphs (KS 19208).

PROG. LINN. SOG. N.S.W., 111 (2), 1989

C.M. A. KENNEDY De

Fig. 2. Conicochernes doyleae sp.nov., scanning clectron micrographs of female paratypes: F, trochanter of right
pedipalp; G, carapace, dorsal aspect (pedipalps dissected); H, pleural membrane.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

126 NEW AUSTRALIAN PSEUDOSCORPION

Diagnosis: A species of the genus Conicochernes with the following characters:
trichobothrium zt widely separated distally from cst; serrula exterior with 16-17 equi-
dimensionally broad, blunt lamellae and a long, narrow acuminate proximal lamella; 2
galea setae present, eye spots distinct.

Description: Adults. Colour dark red-brown, body dorso-ventrally compressed, surface
of carapace predominantly papillate; pleural membrane with closely set undulating
longitudinal folds (Fig. 2H). Pedipalp stout, trochanter with gently rounded anterior
margin, setae very long and acuminate proximally (Fig. 2F), elsewhere short, straight,
longitudinally ribbed in upper distal third and terminating in 3-4 small denticles of
different lengths (Fig. 1C); sculpture coarsely papillate; L/W ratio 1.15-1.40 (0°), 1.20-
1.40 (Q); femur stout, anterior margin relatively straight, posterior margin well
rounded proximally, abruptly pedicellate, setae short, terminally denticulate as
described above, 2.75-3.00 (0°), 2.56-3.00 (Q); tibia stout, anterior margin gently
rounded, distinctly pedicellate 1.69-2.40 (oO), 1.63-2.33 (Q); chela very stout, chelal
hand broad proximally, narrowing distally and longer than chelal fingers, lateral
margins relatively straight, chela (with pedicel) 2.93-3.44 (0), 3.02-3.44 (9), chela
(without pedicel) 2.43-3.17 (O°), 2.75-3.25 (Q ) x longer than broad. .Fixed chelal finger
with 8 trichobothria (Fig. 4M, N), z¢ well separated distally from ist; movable finger with
4 trichobothria, venom tooth large; marginal teeth on each finger 40-50 contiguous,
accessory teeth present; sense spots present; male chelal fingers with extensive medial
gape (Fig. 1A); female chelal fingers without gape (Fig. 1B). Chelicera small with 5
setae, lamina exterior present, serrula exterior attached throughout with 18-19 (0°), 17-
18 (Q ) equal broad, blunt lamellae and a long acuminate proximal lamella (Fig. 1D);
galea well developed, O with 3-5 small rami, 9 5-6 rami, in most cases 2 galea setae
present (Fig. 1E); flagellum with 3 blades of unequal length, distal blade long with 8-9
small spines, approximately 3.40um in length, spaced 4.0-4.70um along the anterior
face (Fig. 1Da). Carapace: anterior margin truncate; pars cephalica covered with trans-
verse discontinuous ridges and corresponding furrows that may be modified by very
small, transversely aligned papillae, the remaining part of carapace coarsely and
densely papillate; posterior margin slightly angulate with centrally elevated disc, lateral
margins slightly converging distally (Fig. 2G); setae short, terminally denticulate, 6: 10:
60-70 (O°), 6: 13-14: 50-60 (9 ); L/W ratio 1.05-1.16 (07), 1.00-1.16 (9) x longer than
broad, eye spots present. Tergites: interscutal membrane in shallow folds, setae short,
terminally denticulate; chaetotaxy: O& 12-14: 14:: 14-16: 14: 16-17: 16-18: 18-20; 18-20: 17-
18: 18-20: 14-16: ?; 9 10-14: 12-15: 16-17: 16-18: 18-20: 18-20: 18-20: 18-20: 18-20: 13-15:
Qe @ ronseroaire, (Coxe cnaciotarays Cr 2; 53 24-25, Os Ss WOR2, Oz ae WD, Oz Ss WO=2, Os 3s
NDAs © Ye Be WO=io, We Ze WB, ie Ze OQ. te Be Jos. We Ze IS, Malle genntalas close
apodemes elongate, tapering; paired lateral apodemes extend ventrally and laterally to
curve upwards terminally (Fig. 3K). Chaetotaxy: anterior operculum large, posterior
margin with row of 7 acuminate setae curling into genital aperture, anterior to these, a
row of 9 short setae above which are 3-4 long setae forming a pyramidal effect and
surrounded by 11-12 very short setae; posterior operculum small with only 10-12
scattered setae (Fig. 31). Female genitalia: with lateral apodemes which provide for
muscle attachment and support of the genital atrium; two median cribriform plates
associated with the medium accessory glands and two lateral cribriform plates on the
anterior wall of the lateral diverticula, each plate cuticular and perforated by pores;
spermathecae two blind, digitate projections resulting from antero-dorsal evagination of
median diverticulum. Scattered cells occur in epithelium of median diverticulum (Fig.
3L). Chaetotaxy: posterior margin of anterior operculum with row of 27-30 acuminate
setae, long over small genital aperture, becoming short laterally, anterior to these and

PROG. LINN. SOG. N.S.W., 111 (2), 1989

C.M. A. KENNEDY 27

0.1mm

Fig. 3. Conicochernes doyleae sp.nov., 1 and J, scanning clectron micrographs of genital region, external aspect,
male and female paratypes respectively. K, male genitalia; L, female genitalia.

centrally situated 6-7 very long setae above which are 7-8 short setae; posterior oper-
culum with 7-10 setae distributed centrally at random, posterior margin with 17-18
uniseriate acuminate setae (Fig. 3J). Sternites: setae short, acuminate; chaetotaxy: O ():
27-30: 12-14: 16-18: 20-23: 18-20: 18-20: 18: 18: 10-12: 10: °?; Q 0: 30-40: 20-24: 16-18: 16-
SOG a loalO-18424-26) 282% stermites4-1l mmisermate:

Dimensions (mm): body length 2.70-3.28 (2.92-3.46); pedipalps: trochanter 0.36-
0437 0227-0234 (0:36-0739/0:27-0:30), femur (027.7-0:90/0-27-0:30 (0:75-0584/0.27-0:30);
tibia 0.59-0.72/0.30-0.39 (0.56-0.66/0.27-0.36), chela (with pedicel) 1.08-1.30/0.34-0.43
(1.18-1.26/0.36-0.41), chela (without pedicel) 1.02-1.24/0.34-0.43 (1.08-1.17/0.36-0.41),
movable finger length 0.46-0.54 (0.43-0.56); chelicera 0.18-0.23/0.10-0.14 (0.21-0.25/0.11-
0.14) movable finger length 0.12-0.16 (0.12-0.18); carapace 0.84-0.98/0.75-0.89 (0.86-
1.03/0.84-0.98); cucullus length 0.14-0.19 (0.16-0.22) ocular breadth 0.56-0.64 (0.56-
0.64); leg I: coxa width 0.30-0.36 (0.32-0.36), trochanter 0.13-0.16/0.11-0.16 (0.14-
0.16/0.11-0.16), basifemur 0.16-0.22/0.14-0.18 (0.16-0.25/0.14-0.17), telofemur 0.22-
0.33/0.13-0.16 (0.22-0.33/0.12-0.16), tibia 0.16-0.28/0.10-0.14 (0.22-0.28/0.10-0.14), tarsus
0.16-0.22/0.08-0.10 (0.16-0.22/0.07-0.08); leg 4: coxa width 0.28-0.33 (0.30-0.36),
trochanter 0.18-0.25/0.14-0.16 (0.19-0.28/0.14-0.19), basifemur 0.25-0.30/0.16-0.25 (0.19-
0.28/0.16-0.22), telotemur 0.39-0.50/0.19-0.24 (0.41-0.50/0.21-0.28), tibia 0.28-0.42/0.12-
0.14 (0.28-0.40/0.11-0.15), tarsus 0.22-0.28/0.08-0.10 (0.16-0.20/0.07-0.11).

Tritonymph. L/W ratio: pedipalpal trochanter 1.36-1.73, femur 2.12-3.21, tibia 1.56-
2.05, chela (with pedicel) 3.00-3.65, chela (without pedicel) 2.78-3.30 x longer than
broad. Fixed finger with 7 trichobothria, movable finger with 3 trichobothria, sb and ist
absent; serrula exterior of chelicera with 13 broad lamellae, distal and proximal lamellae
long and narrow. Carapace: posterior margin angulate, 6: 10: (50-60) setae, 1.04-1.20 x

PROG. LINN. SOG. NSW... 111 (2). 1989

128 NEW AUSTRALIAN PSEUDOSCOR PION

Fig. 4. Conicochernes doyleae sp.nov., female paratypes: M and N, scanning electron micrographs of right chela,
lateral, external and internal aspects respectively showing position of trichobothria; P, Ist left leg: Q 4th left
leg; R, left pedipalp, dorsal.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

C.M. A. KENNEDY 129

longer than broad. Tergal chaetotaxy: 10-12: 12: 12: 12: 13-14: 11-12: 14-15: 14-15: 12: 10-
12: 12-14: 2. Sternal chaetotaxy: 0: 6-8: 8-10: 12: 14-16: 16: 16-18: 10-14: 12-18: 10-12: 12-
aC oxalehactotaxy. O2e44 Oi2 7/0 0250-7)) 02 wO=I3y Nlonotarsate:

Dimensions (mm): body length 2.38-2.88; pedipalps: trochanter 0.26-0.33/0.19-0.22,
femur 0.47-0.61/0.19-0.25, tibia 0.39-0.47/0.19-0.25, chela (with pedicel) 0.84-0.95/0.25-
0.30, chela (without pedicel) 0.78-0.86/0.25-0.30, movable finger length 0.35-0.45;
carapace 0.61-0.70/0.56-0.64.

Deutonymph. L/W ratio: pedipalpal trochanter 1.35-1.72, femur 2.14-2.57, tibia 1.56-
2.00, chela (with pedicel) 3.19-4.06, chela (without pedicel) 2.76-3.68 x longer than
broad. Fixed finger with 6 trichobothria, movable finger with 2 trichobothria, st, sb, st,
esb absent, serrula exterior of chelicera with 11 blunt lamellae, distal and proximal
lamellae long and narrow. Carapace 6: 8: (30-40) setae, 1.00-1.07 x longer than broad.
Wergal chaetotaxy: ‘8: 8: 8:-1012: 12: 10: 10-12: 12; 12: 10-112: 8: 2. Sternal chaetotaxy:
0: 0: 0: 4-6: 6-10: 7: 8-10: 9-11: 10-12: 9-10: 10: 1. Coxal chaetotaxy: 0: 2-3: 6-7, 0: 3: 5-6,
9227902937).

Dimensions (mm): body length 1.48-2.24; pedipalps: trochanter 0.19-0.25/0.11-0.16,
femur 0.30-0.42/0.14-0.19, tibia 0.25-0.33/0.12-0.19, chela (with pedicel) 0.61-0.73/0.16-
0.22, chela (without pedicel) 0.56-0.65/0.16-0.22, movable finger length 0.25-0.31;
carapace 0).39-0.47/0.37-0.45.

Protonymph. L/W ratio: pedipalpal trochanter 1.20-1.33, femur 1.75-2.50, tibia 1.58-
1.91, chela (with pedicel) 3.14-4.30, chela (without pedicel) 3.00-3.90 x longer than
broad. Fixed finger with 3 trichobothria, ¢, et, eb, isb present. Serrula exterior of chelicera
with 10 broad lamellae, distal and proximal lamellae long and narrow. Carapace with 6:
7-8: (25-30) setae, 1.04-1.10 x longer than broad. ‘Tergal chaetotaxy: 6: 8: 6: 6: 6: 8: 6-8:
OCHO an otehnalachiactotaxyai0- 02-3214 40410-0110 0450-G410-6410-O510 1051-25 @.oxall
Ghactoraxy 102410 geo O30 Ze 3.

Dimensions (mm): body length 1.10-1.45; pedipalps: trochanter 0.12-0.16/0.10-0.13,
femur 0.19-0.28/0.10-0.12, tibia 0.18-0.23/0.10-0.12, chela (with pedicel) 0.43-0.54/0.12-
0.14, chela (without pedicel) 0.39-0.48/0.12-0.14, movable finger length 0.18-0.21;
carapace 0.28-0.48/0.26-0.43.

ACKNOWLEDGEMENTS
I wish to express my thanks to Dr P. D. Hillyard (British Museum (Natural
History) ) for the loan of type material and to Dr M. Gray (Australian Museum) for
reviewing the manuscript and offering comments thereon.

References

Brier, M., 1948. — Uber Pseudoscorpione der Australischen Region. Eos Madr. 24: 232-536.

CHAMBERLIN, J. C., 1931. — The arachnid order Cheloncthida. Stanford Univ. Publy. (Biol) 7: 1-284.

Harvey, M. S., 1981. — A checklist of the Australian Pscudoscorpionida. Bull. Br. arachnol. Soc. 5 (5):
ZIP 2S

——,, 1985. — Pscudoscorpionida. Jn D. W. WALTON, (ed.), Zoological Catalogue of Australia, 3: 133. Canberra:
Australian Government Publishing Service.

LEGG, G., 1974a. — A gencralised account of the female genitalia and associated glands of pscudoscorpions
(Arachnida). Bull. Br. arachnol. Soc. 3: 42-48.
——, 1974b. — A generalised account of the male genitalia and associated glands of pscudoscorpions

(Arachnida). Bull. Br. arachnol. Soc. 3: 66-74.

PROG. LINN. SOG. N.S.W,, 111 (2), 1989

oe
‘yh
vit

rie, i ee f
wr ware ‘
a _flastiied |

! ‘

Pic aay

“ Wal te

Mckenziartia and FPectocythere

(Pectocytheridae, Ostracoda, Crustacea) in
Lake Macquarie, New South Wales

IRADJ YASSINI and MARY MIKULANDRA

YASSINI, I., & MIKULANDRA, M. Mckenzzartia and Pectocythere (Pectocytheridac,
Ostracoda, Crustacea) in Lake Macquarie, New South Wales. Proc. Linn. Soc
N.S.W, 111 (2), 1989: 131-139)

The distribution pattern of four species of Mckenziartia and one species of
Pectocythere in Lake Macquaric, New South Wales, is investigated and three new species:
Mckenziartia thomi sp. nov., Mckenziartia mowbrayi sp. nov. and Fectocythere royi sp. nov., arc
described.

Trad) Yassini, Dept of Geology, University of Wollongong, Wollongong, Australia 2500, and Mary

Mikulandra, Dept of Geography, University of Sydney, Australia 2006; manuscript received 26 July

1988, accepted for publication 15 February 1989.

INTRODUCTION

The genus Mckenziartia was described by Bentley (1988) on the basis of carapace
morphology. The genus was erected to designate a group of pectocytherid ostracodes
characterized by the presence of a crescentic fossa over the central muscle field, a large
anterior vestibulum with short, simple or occasionally-branched marginal pore canals,
and an antimerodont hinge structure. The crescentic fossa over the central muscle field
was described as a diagnostic feature of the genus. Examination of a large population of
the species from different estuaries of New South Wales shows, however, that the size
and shape of fossa may vary and is often integrated with the ornamentation pattern of
the valves. The size and the shape of the vestibulum and the distribution of the anterior
marginal pore canals remain unchanged, and they present a more stable diagnostic
feature for the genus.

Mckenziartia Bentley is a typical inhabitant of the estuarine environment and has
been reported from several estuaries around Australia (Bentley, 1988).

Present work investigated the distribution pattern of four species of Mckenzzartia:
Mckenziartia portjacksonensis (McKenzie, 1967), Mckenziartia foveata (Hartmann, 1978),
Mckenziartia thomi sp. nov. and Mckenziartia mowbrayt sp. nov., as well as distribution of
Fectocythere royt sp. nov., another member of the Pectocytheridae in Lake Macquarie. The
type specimens are deposited at the Australian Museum. Some paratype specimens
were also deposited at the Commonwealth Palaeontological collection in the Bureau of
Mineral Resources, Canberra.

AREA STUDIED

Lake Macquarie is located some 85km north of Sydney, on the central coast of New
South Wales. With an extremely irregular outline, the lake extends in a north-south
direction for over 22km, has a width of 10km and a surface area of 110km?.

The average depth of the lake is 6.7m and its maximum depth never exceeds 12m.
A narrow inlet channel 3.5km in length and 240m to 400m wide connects the lake to the
Pacific Ocean.

The lake has a total catchment area of 770km*. Three major creeks; Mannering
Creek in the south, Dora Creek in the west and Cockle Creek in the north, contribute up
to 4% of the total lake volume (Bass Becking et al., 1959). Some seven other minor creeks

PROG. LINN. SOG. N.S.W., 111 (2), 1989

132 OSTRACODES FROM LAKE MACQUARIE

pew CLD. pn
Sik, ARGS

Lake Macquarie
Kel

Built-up areas

@ Sample location

Vv Sample with
Mckenziartia

x No ostracodes

Fig. I, Location map of the samples studied.

PROC. LINN. SOC, N.S.W., 111 (2), 1989

I. YASSINIAND M. MIKULANDRA 133

TABLE 1

Species and number of valves in 30m of samples

; : Samples

pipes BO | A ol SOP, Se VE ONE TT AE GE 8
Mckenziartia portjacksonensts 2... ... 26 10 245 Uae DES 8 ils} 40) 14 3 2
Mlekemznamtina WHORTD o555705000000¢ — — 48 = = = = pas aa = = =
Mckenziartia foveata ............. _ — 12 — — = = = ae = = -
Mckenziartia mowbrayt ........... = = D) =: a = ls 3 zs =e 8

[Pace MHORE THOME sooo adoacee doce. = —

also flow into the lake during the rainy periods. A total of 26 grab samples were taken
from the bottom sediment of the lake; Fig. 1 indicates the location of the samples
studied. Table 1 shows the number of valves of each species in 30ml of unwashed sample.

SYSTEMATICS
Class OSTRACODA Latreille, 1802
Order PODOCOPIDA Muller, 1894
Family PECVOCYTHERIDAE Hanai, 1957
Genus Mckenziartia Bentley, 1988
MCKENZIARTIA PORTJACKSONENSIS (McKenzie, 1967)
Fig. 2A-D and Fig. 4A

‘Hemicytheridea portjacksonensis McKenzie, 1967: p. 85, pl. 12, figs 31-j,6.
Munseyella’ tumida Swanson, 1979: p. 158, fig. 4q.
Fectocythere portjacksonensis (McKenzie), Hartmann, 1980: p. 122, pl. 5 fig. 17.
FPectocythere portjacksonensis (McKenzie), Yassini and Jones, 1987: p. 826, pl. 2, fig. 21.
Mckenziartia portjacksonensis (McKenzie), Bentley, 1988: p. 445, pl. 1, figs. e-f, text fig.
4a-b.

Distribution: The species has been recorded from many estuaries and shallow open
marine environments around Australia (McKenzie, 1967; Hartmann, 1978, 1980;
Yassini and Jones, 1987; Bentley, 1988) and from New Zealand (Swanson, 1979). In
Lake Macquarie, it was recorded from 12 stations but it was particularly abundant at
station 47 at the mouth of the inlet channel (Table 1).

Mckenziartia foveata (Hartmann, 1978)
Fig. 2G-I, Fig. 3G and Fig. 4B
FPectocythere foveata Hartmann, 1978: p. 144, pl. 14, figs. 12-13.
Fectocythere foveata Hartmann, Yassini and Jones, 1987: p. 826, pl. 2, figs. 18-20.

Remarks: The hinge structure and vestibulum pattern justify attribution to the genus
Mckenziartia.

Distribution: The species has been recorded from several estuaries and shallow open
marine environments around the continent (Hartmann, 1978; Yassini and Jones, 1987).
In Lake Macquarie it was only found in sample 47, dredged from a seagrass bed close to
the inlet channel (Table 1).

Mckenziartia thomi sp. nov.
Fig. 3A-F and Fig. 4C

Diagnosis: Carapace elongate, length about 2.6 times height. Anterior broadly
rounded, posterior rounded ventrally, subvertical medio-dorsally. Dorsum straight,

PROG. LINN. SOG. N.S.W., 111 (2), 1989

154 OSTRACODES FROM LAKE MACQUARIE

Fig. 2. A-D, Mckenziartia portjacksonensis (McKenzic): A, RV, ©, external view; B, 6, LV, internal view; C, 9,
ventral view of the carapace; D, 6, dorsal view of the carapace, sample 47. E, F, J-L, Mckenziartia mowbrayi sp.
noy.; E, LV, 9, external vicw, holotype AM P37639; F, RV, 6, external view, paratype AM P37641; J, 9,
ventral view of the carapace, paratype AM P37641; K, 9, dorsal view of the carapace, paratype AM P37642;
L, LV, 6, internal view, paratype AM P37642, sample 1. G-I, Mckenziartia foveata (Hartmann); G, RV, 6,
external view; H, 6, dorsal view of the carapace; I, LV, 6, internal view, sample 47.

ventrum gently concave. Valves equal in size. In dorsal and ventral views, carapace elon-
gate with parallel borders. Internally: inner lamella broad, anterior vestibulum deep,
narrow, posterior vestibulum broad and elongate. Marginal pore canals simple, few and

PROG. LINN. SOG. N.S.W., 111 (2), 1989

I. YASSINI AND M. MIKULANDRA 1335)

Fig. 3. A-F, Mckenziartia thomi sp. nov.; A, LV, O,, external view paratype AM P37647; B, LV, 9, internal view,
paratypc AM P37648; D, RV, 9, external view, holotype AM P37646; C, RV, oO’, internal view, paratype AM
P37649; E, oO’, ventral view of the carapace, paratype AM P37650; F, o&, dorsal view of the carapace, para-
type CPC 26554, sample 47. G, Mckenziartia foveata (Hartmann) ©’, ventral view of the carapace, sample 47.
H-K, Pectocythere royi sp. nov.; H, RV, internal view, holotype AM P37661; I, LY, internal view, paratype AM
P37662; J, dorsal view of the carapacc, paratype AM P37663; K, ventral view of the carapace, paratype AM
P37664, sample 47.

scattered. Hinge antimerodont. Muscle scars: four adductor scars in a subvertical row,
two frontal scars and a large furcal scar. Valve surface reticulated, with three distinct

PROC. LINN. SOG. N.S.W., 111 (2), 1989

136 OSTRACODES FROM LAKE MACQUARIE

concentric ridges parallel to posterior margin and a set of oblique ridges joining the
ventral ridge.

Remarks: With a crenulate posterior socket, and a straight and crenulate median bar
the species clearly belongs to the Pectocytheridae and it can be differentiated from the
allied species (Mckenziartia portjacksonensis (McKenzie) and Mckenziartia mowbrayi sp. nov.)
in the details of surface ornamentation.

Material: Holotype AM P37646, right valve, adult male; length 0.51!mm. Paratypes
AM P37647, AM P37648, AM P37649, AM P37650 and CPC 26554. All from type
locality.

Type locality: Lake Macquarie entrance tidal channel, New South Wales, Australia
(lat. 33°03’00"S; long. 151°38’00”E); Recent, estuarine, muddy sand, with Zostera
capricornt (Aschers.); depth 5.2m; on the sampling day salinity was 35.8°/o0, water
temperature 22.8°C, and dissolved oxygen 6.6mg/I.

Etymology: The species is named after Professor Bruce Thom, Geography Depart-
ment, University of Sydney.

Distribution: The species has also been recorded from Heron Island, off Queensland
(Labutis, 1977) and Broken Bay, New South Wales (Yassini, unpublished).

Mckenziartia mowbrayi sp. nov.

Fig. 2E-F, J-L and Fig. 4D

Diagnosis: carapace elongate, length about 2.3 times height. Anterior broadly rounded,
posterior rounded to gently subacute. Dorsum straight, ventrum gently concave in the
middle. Valves equal in size. In dorsal and ventral views, sides of the carapace are
parallel. Internally: inner lamella broad, anterior vestibulum shallow and broad.
Posterior vestibulum narrow. Marginal pore canals straight, well-spaced and few. Hinge
antimerodont. Valve surface reticulate, with a few distinct, concentric low ridges at the
posterior end, and three at the anterior end. In ventral view, the ridges form a series of
chevron-type grooves converged toward anterior and posterior ends. Muscle scars are
typical of the genus Mckenzzartia (Bentley, 1988).

Remarks: When compared with Mckenzzartia thomi sp. nov, and Mckenziartia portjacksonen-
sis (McKenzie), it differs in its typical surface reticulation.

Material: Holotype AM P37639, carapace, male; length 0.46mm. Paratypes AM
P37640, AM P37641, AM P37642, AM P37643 and CPC 26553. All from type locality.

Type locality: Lake Macquarie (Sample 1), north of Mannering Park, New South
Wales, (lat. 33°09'00"S; long. 151°33’00”E); Recent estuarine, sandy mud; depth
3.7m; on the sampling day salinity was 36.22°/o0, water temperature 22.8°C, and
dissolved oxygen 4.4mg/1.

Etymology: After Mr William Mowbray, Chairman of Lake Illawarra Management
Committee.

Distribution: Outside Lake Macquarie, the species has also been found in Broken Bay
(Yassini, unpublished) and off Queensland (Labutis, 1977).

PROC. LINN. SOG. N.S.W., 111 (2), 1989

I. YASSINIAND M. MIKULANDRA 13

100pm.

Fig. 4. A, Mckenziartia portacksonensis (McKenzic), RV, internal structure; B, Mckenzvartia foveata (Hartmann),
RV, internal structure; C, Mckenziartia thomi sp. nov., RV, internal structure, holotype AM P37546; D,
Mckenztartia mowbrayi sp. nov., RV, internal structure, paratype AM P37644; E, Pectocythere royt sp. nov., LV,
internal structure, paratype AM P37638.

PROG, LINN. SOG. N.S.W., 111 (2), 1989

158 OSTRACODES FROM LAKE MACQUARIE

Genus Pectocythere Hanai, 1957
Pectocythere royi sp. nov.
Fig. 3H-K and Fig. 4E
Fectocythere sp. (Ceduna 120), Hartmann; (Nec Hartmann, 1980) Yassini and Jones, 1987;
Oe BAG, OL 2, ine, Weeclk/,

Diagnosis: Carapace elongate, small, length about 2.1 times height. Anterior broadly
rounded, posterior with an elevated rim and _ postero-ventrally oblique. Dorsum
straight, slightly inclined posteriorly. Ventrum straight, valves ornamented with irregu-
larly distributed rounded or elongated alveoh. In dorsal view, broader posteriorly. Inter-
nally: inner lamella broad, anterior vestibulum broad and shallow, marginal pore canals
few and straight. Hinge merodont, consisting in the right valve of an elongate anterior
tooth, distinctly crenulated median groove and coarsely lobated posterior terminal
tooth. Central muscle scars comprising a row of four adductors and a V-shaped antennal
scar. Sex dimorphism is not known.

Remarks: The only described species with closely similar shape is Pectocythere sp.
(Ceduna 120) Hartmann (1980) which 1s easily distinguished from the present species by
the depression of the postero-ventral region and lack of the raised posterior rim.

Material: Holotype AM P37661, right valve: 0.43mm length. Paratype AM P37662,
AM P37663, AM P37664 and CPC 26555. All from type locality.

Type locality: Lake Macquarie (Sample 47), entrance tidal channel, New South Wales
(lat. 33°03°00"S, long. 151°38’°00”E); Recent, estuarine, muddy sand with Zostera
capricorni (Aschers.); depth 5.2m; on the sampling day, salinity was 35.8°/o0, tempera-
ture 22.8°C and dissolved oxygen 6.6mg/I.

Etymology: After Dr Peter Roy, Geological Survey, New South Wales.

Distribution: Widely distributed in estuarine environments along the coast of New

South Wales.

DISCUSSION

Among the five described species, Mckenzzartia portjacksonensis is the only one which
is widespread and abundant in all the estuarine environments along the eastern coast of
Australia; from Heron Island in the north (Labutis, 1977) to Bass Strait in the south
(Yassini and Blom, in preparation). Pectocythere roy1, has also been found in limited
numbers in Broken Bay, Twofold Bay and Bass Strait. Mckenzzartia foveata, initially
described by Hartmann (1978) from Port Hedland, Western Australia, was also found in
several estuaries of New South Wales (Lake Illawarra, Botany Bay, Broken Bay,
Brisbane Waters). The species was also reported by Labutis (1977) from the Great
Barrier Reef, off Queensland. The other two species; Mckenzzartia thomi sp. nov. and
Mckenziartia mowbrayi sp. nov. appear to be limited to the northern (Labutis, 1977) and
central coast of New South Wales.

References
Bass BECKING, L. G. M., THOMSON, J. M., and Woon, F. J., 1959. — Some aspects of the ecology of Lake
Macquarie NSW with regard to alleged depletion of fish. I. General introduction. Aust. J. Mar, Fresh-
water, Rey. 10; 297-305.
BENTLEY, C., 1988. — Podocopida Ostracoda of Brisbanc Water, near Sydney, south-castern Australia. JT.
HANAt, N. IKeYA and K. ISHIZAKI, (eds), Evolutionary Brology of Ostracoda: 434-448. Amsterdam:
Elsevier.

PROG. LINN. SOG. N.S.W., 111 (2), 1989

I. YASSINIAND M. MIKULANDRA 159

Hanal, T., 1957. — Studies on the Ostracoda from Japan. 1 — Subfamily Leptocytherinac n. subfam. /. Fac
Sci. Univ. Tokyo Sect. 11 10: 469-82.
HARtMANN, G., 1978. — Die Ostracoden der Ordnung Podocopida G. W. Muller, 1894 der tropisch-

subtropischen Westkuste Australicns (zwischen Derby im Norden und Perth in Suden). Afitt, Ham
Zool. Mus. Inst. 75: 53-219.

——., 1980. — Dic Ostracoden der Ordnung Podocopida G. W. Miller, 1894 der warmtempericrten und
subtropischtropischen kustenabschnitte der Sud-und Sudostkuste Australiens (zwischen Ceduna im
Westen und Lake Entrance im Osten). Mitt, Ham. Zool. Mus. Inst. 77: V1-204.

Laputis, V. R., 1977. = Cytheraccan Ostracoda from the Great Barricr Reef. North Ryde, N.S.W.: Mac-
quaric University, M.Sc. (Hons) thesis, unpubl.

LArREILLE, P. A., 1802. — Histoire des Cypris et des Cytheres. In, LAPREILLE, P. A., (ed.), Histoire Naturelle
Generale et Particulvere des Crustaces et des Insectes. Vol. 4: 232-254. Paris: Dulart.

McKenzir, K., 1967. — Recent Ostracoda from Port Phillip Bay, Victoria. Proc Roy. Soc. Victoria 80: 61-106.

MULLER, G. W., 1894. — Die Ostracoden des Golfes von Neapel und der angrenzenden Meeres-abschnitte.
Naples Sta. Zool. Fauna Flora Golfes Neapel, Monographie, 31: 1-404.
SWANSON, K. M., 1979. — Recent Ostracoda from Port Pegasus, Stewart Island, New Zealand. New Zealand
J. Mar. Freshwater Research 13: 151-170.
YASSINI, I., (in prep.). — Recent shallow water estuarine ostracods from New South Wales.
,and Bom, W., (in prep.). — Distribution of Recent ostracods in Bass Strait, south east Australia.
, and Jones, B. G., 1987. — Ostracoda in Lake Illawarra, environmental factors, assemblages and

systematics. Aust, J. Mar. Fresh. Research 38: 795-845.

PROG. LINN. SOG. N.S.W., 111 (2). 1989

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PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 111
NUMBER 3

Wetlands of the Lower Clarence Floodplain,
Northern Coastal New South Wales

IRS I, JPRIBSSIENZ
(Communicated by P. ADAM)

PRESSEY, R. L. Wetlands of the lower Clarence floodplain, northern coastal New South
Wales. Proc. Linn. Soc. N.S.W. 111 (3), 1989: 143-155.

The wetlands on the lower Clarence floodplain, on the far north coast of New
South Wales, were mapped and described during a detailed survey in 1982. The survey
identified 755 wetlands with a total area of about 14,700 ha. Most wetlands were smaller
than 10 ha but the relatively few large wetlands made up most of the total area. Open
water occupied only 5% of the total wetland area at the time of the survey although
some open water occurred in 45% of wetlands. The most important plant families, in
terms of extent and frequency of occurrence, were Poaceae (covering 23% of the wet-
land area, occupying 82% of the total number of wetlands), Cyperaceae (20% area,
36% number), Myrtaceae (17% area, 36% number), Polygonaceae (12% area, 76%
number) and Juncaceae (10% area, 75% number). Most wetlands were fringed by
dense stands of herbaceous plants but, due to clearing, had few trees on their margins.
Ninety-eight percent of wetlands were grazed to some extent and 92% of the total wet-
land area was affected by drainage. Most wetlands are seasonal with relatively small
catchments. Only 28% had catchments on the bedrock slopes surrounding the flood-
plain. Most catchments have been completely cleared for agriculture. Most wetlands
are freehold tenure.

R. L. Pressey, New South Wales National Parks and Wildlife Service, PO. Box 1967, Hurstville,
Australia 2220; manuscript received 1 November 1988, accepted for publication 15 March 1989.

KEY WORDS: Wetlands, survey, plant taxa, catchments, Clarence floodplain.

INTRODUCTION

The coastal wetlands of New South Wales have been mapped and classified in a
number of broad-scale surveys. Hannah’s (1968) landscape approach dealt mainly with
the wetlands of the dune systems and was restricted to the north coast. West et al., (1985)
mapped and classified the vegetation of the State’s estuarine wetlands. The surveys of
Goodrick (1970) and the Coastal Council of New South Wales (1985) were of dunal, estu-
arine and floodplain wetlands along the whole coast. The Coastal Council study simply
delineated wetlands for the purposes of development controls under State Environmen-
tal Planning Policy No. 14. Goodrick’s survey provided a useful classification based on
dominant vegetation and water regime and is the best overview of wetlands on the
coastal floodplains. It does not, however, provide sufficiently detailed information on
individual floodplain wetlands for conservation or management.

On the Clarence floodplain, none of the previous local, more detailed studies of
flora and fauna has covered the wetlands comprehensively. They have concerned only
one or a few wetlands and have had a variety of purposes and approaches. For example,
Broome (1978) surveyed waterbirds in several areas and two environmental impact
statements (Clarence River County Council, 1978, 1980) include vegetation maps and
descriptions of some large wetlands. Most other publications on the Clarence floodplain
wetlands deal with soils and specific occurrences of birds (Pressey, 1981).

Wetland surveys for the National Parks and Wildlife Service, designed to be both
detailed and comprehensive, covered the lower floodplains of the Hunter, Clarence and
Macleay Rivers in 1981, 1982 and 1984, respectively, and the coastal lowlands of Tweed
Shire in 1986.

The main aims of these surveys were to: plot wetlands with accuracy and, in the

PROC. LINN. SOC. N.S.W., 111 (3), 1989

144 LOWER CLARENCE WETLANDS

Tweed survey, associated coastal communities on 1:25 000 topo-cadastral maps;
describe the wetlands and their catchments from features that could be recorded during
a single field visit or remotely from maps and aerial photographs; rank the wetlands for
nature conservation, based on a combination of selected criteria.

This paper summarizes the results of the inventory and description of wetlands on
the lower Clarence floodplain. Full data, the results of conservation evaluation, and
reduced wetland maps have all been incorporated in a consultancy report (Pressey,

1987).

STUDY AREA

The Clarence is one of the major coastal rivers in New South Wales and has an
extensive floodplain, some 500 km north-east of Sydney (Fig. 1). The lower Clarence
floodplain was defined as alluvium below the 10 m contour, a broad expanse of flood-
prone land downstream of the very narrow, upper floodplain. Virtually all of the alluvial
wetland area in the Clarence system occurs on the lower floodplain. The survey was also
restricted to non-estuarine wetlands (those without saltmarsh and mangroves), although
it included brackish areas intermediate between fresh and full tidal waters.

The study area occupies approximately 700 sq. km (Fig. 1). To the north of the
river, 1t extends upstream as far as Whiteman Creek and includes extensive alluvial flats
to the north of Grafton, south of Lawrence and around the Broadwater. The major wet-
land on this side of the river is the Everlasting Swamp. South of the river the survey
covered areas downstream of Seelands including large expanses of floodplain between
Swan Creek and Tyndale and around Wooloweyah Lagoon. The main wetlands in these
areas are the Harrington Lagoon complex, the heavily wooded Shark Swamp and a
group of large wetlands on the Coldstream River, one of the Clarence’s lower tributaries.
The survey also covered Woodford, Harwood and Chatsworth Islands which lie either in
the course of the river or in its broad estuary.

Virtually all of the floodplain is cleared for farming. The major agricultural use 1s
grazing of beef and dairy cattle. Sugar cane is an important crop in the eastern parts and
relatively small areas are planted with poplars, vegetables and other crops. Drainage of
wetlands began soon after European settlement. Around the turn of the century,
drainage unions or trusts were formed to enable adjacent landholders to arrange for
drainage in co-ordinated systems, with design and construction provided by the Depart-
ment of Public Works. This work was to promote the growth of vegetation considered
valuable as pasture and to encourage closer settlement in the region. From the late 1950s
to the 1970s, under flood mitigation schemes, many drains, levees and floodgates were
constructed and integrated to exclude floods from certain areas and to accelerate the
recession of floodwaters. The alteration of wetlands continued during this period.

Goodrick (1970) found that about 47% of all wetlands in the far north coast region
had been destroyed or significantly altered by 1969, mainly by drainage and flood miti-
gation. About 85% of the total affected area consisted of shallow floodplain wetlands,
with more wetland now affected since flood mitigation on the Clarence has continued
since Goodrick’s survey (Pressey and Middleton, 1982). In addition, many private
developments have recently drained and cleared wetlands.

METHODS

The units of the survey were discrete wetland basins which were initially identified
on black and white aerial photographs as areas of darker tone or as stands of trees. ‘The
available aerial photographs were taken between August 1978 and March 1979 and
varied in scale from 1:25 000 to 1:40 000. The minimum size for recognition of wetlands

PROG. LINN. SOC. N.S.W., 111 (3), 1989

Re Ly PRESSEY 145

<— STUDY AREA

Whiteman
Creek 4

Coldstream
River

x. It
SOUTH GRAFTON

Fig. I. The study area. Numbers are those given to named wetlands during the survey. The names corres-
ponding to these numbers are: 9 Double Swamp; 11 Bunyip Creek; 31 Alumy Creek; 54 Southgate Lagoon; 61
Southgate Creek; 62 Franks Creek; 72 Long Waterhole; 84 Hannons Lagoon; 90 Harrisons Creek/Coxs
Creek; 123 Everlasting Swamp; 172 Mororo Creek; 181 Duckpond Creek; 206 Elbow Creek; 210 Roberts
Creek; 257 Cowans Ponds; 278 Cassons Creek/Harrington Lagoon/McLachlan Waterhole; 302 Swan Creek;
306 Ellis Swamp/Crowsnest Swamp; 308 Horseshoe Waterhole; 400 Morans Swamp/Colletts Swamp; 413
Chaffin Swamp; 429 Champions Swamp; 436 Stokes Waterhole; 437 Stokes Waterhole; 464 Sweeneys Creek;
496 Sweeneys Swamp/Oregon Creek; 514 McPhees Swamp; 515 McPhees Swamp; 527 Calliope Creek; 573
Calligans Creek; 638 Alumny Creek; 649 Swampy Creek; 703 Shark Swamp/Tyndale Swamp.

was about 0.1 ha. A few small wetlands of 1 ha or less were omitted because of difficult
access.

Final boundaries of wetlands were marked on aerial photographs during a single
field visit to each one, either in summer (3.2.82-28.2.82) or winter (19.7.82-17.8.82).
These boundaries were later transferred onto 1:25 000 topo-cadastral maps. Scale
differences and lens distortion were corrected in the transfer by enlarging or reducing
the images on the photographs and by matching features such as fencelines, roads and
drains between photographs and maps over small areas.

The number and types of attributes recorded for each wetland were constrained by
the time available. In particular, fauna could not be described reliably, and faunal

PROG. LINN. SOG. N.S.W,, 111 (3), 1989

146 LOWER CLARENCE WETLANDS

habitat and importance in each wetland were generally inferred from open water and
the type and structure of vegetation.

Of the twelve attributes listed for each wetland, five were used solely for ranking the
wetlands for conservation (see Pressey, 1987). The remaining seven were used descrip-
tively and are as follows.

1) Szze

Size of wetlands was estimated from corrected boundaries on topo-cadastral maps

using a dot grid.

i) Vegetation/habitats

Vegetation and faunal habitat of the wetlands were described by plant species or
genera and two categories of open water: deep (>60 cm) and shallow <60 cm).
Authorities for plant names are those in Jacobs and Pickard (1981) as amended by Jacobs
and Lapinpuro (1986), unless indicated otherwise. Water depth was judged from fence-
lines and from the slope of the bed. Submerged plants were not recorded, nor were
species from the families Azollaceae or Lemnaceae. The percentage of each wetland
occupied by each taxon or open water category was estimated by eye in the field. Percen-
tages were later converted to absolute areas when the sizes of wetlands were measured.
‘Taxa or open water categories occupying less than 0.1 ha in a wetland were listed only as
present.

Small wetlands were described by walking through or around them, large swamps
by walking through them, by scanning with binoculars from vantage points, and by
visiting areas with distinctive tone or texture on aerial photographs. Some inconsisten-
cies in description would have resulted from differences in sampling intensity between
wetlands. The accuracy of estimates of the percentages of wetland areas occupied by
plant taxa and open water categories would probably be inversely related to wetland
s1Ze.

11) Marginal vegetation

Vegetation on wetland margins was rated in the field according to the percentage of
perimeter lined with trees or emergents (rushes, tall herbs or grasses) and the average
density and width of the fringing emergent band. Fringing plants not typical of wet-
lands, such as tall pasture grasses, were not taken into account as emergents.

iv) Alteration
Signs of obvious alteration such as grazing, drainage and impoundment were
recorded in the field and from aerial photographs.

v) Catchment areas

Wetland catchments were categorized according to whether they were completely
on alluvial flats or at least partly on bedrock slopes. Catchments on bedrock above the 10
m (lowest) contour were delineated and their areas measured on topographic maps.

v1) Catchment land use

Broad types of land use in wetland catchments were recorded in the field and from
maps and aerial photographs. They were listed for each catchment in order of the
proportion occupied. Categories recognized were agriculture (including grazing and
cropping), forestry, industrial areas, mining (only surface extraction of sand, gravel etc.
in the case of the Clarence survey area), natural vegetation, urban areas and waste
disposal (including sewage treatment works).

vil) Ténure
Land tenure, easements and any other relevant land designations were listed for
each wetland in order of the proportion of the wetland affected.

PROG. LINN. SOC. N.S.W., 111 (3), 1989

R. L. PRESSEY 147

RESULTS AND DISCUSSION

The survey covered 755 wetlands with a total area of 14,728 ha. The majority of
wetlands are small (Table 1), 88% of them 10 ha or less in size. These small wetlands
account for only 9% of the total wetland area. The bulk of the wetland area (77%) is
made up by the relatively small number of wetlands (3% of total number) greater than
100 ha in size.

TABLE 1
Size distribution of wetlands
No. of % Total Wetland % Votal
Area (ha) wetlands no. area (ha) area
0-1.0 343 45.4 1749) 2
1.1-5.0 255 33.8 626.6 4.3
5.1-10.0 69 9,1 480.4 Boo
10.1-50 57 7.6 1155.4 7.8
51-100 12 1.6 878 6.0
101-500 13 tod 2713 18.4
501-1000 1 0.1 520 3.0)
> 1000 5 0.7 8180 39).8)
755 100.0 14,7279) 100.0

Seventy-three plant taxa were systematically recorded. Their occurrence in the
study area, with that of the two recorded categories of open water, is summarized in the
Appendix. Some plants were identified only to genus. The most common of these were:

— Juncus spp. refers to species in the Section Genuini, the relative areas of which
were not determined. The only two species identified were /. usztatus and J. poly-
anthemus. From subsequent work on the Macleay floodplain, it is likely that
hybrids of these species are common and widespread.

— Perswaria spp. were not recorded individually. Species collected during the
survey were P. hydropiper (most extensive), P lapathifolia, P. strigosa and F. sp. B.
Another species occurring in the area but not found in the wetlands surveyed is
I& SO, A\

— Typha spp. were also combined. Both 7! domingensis and T: orientalis occur in the
area, with the latter species by far the more common.

Open water was relatively unimportant in terms of the overall area occupied
(Appendix). Shallow open water was more extensive and more commonly recorded than
deep open water. Goodrick (1970) recorded a relatively small area of wetlands domi-
nated by open water on the far north coast generally. The floodplain wetlands in this
region, including those of the Clarence, are largely in the five categories of Goodrick’s
classification that are dominated by herbaceous emergents or by trees: fresh meadows,
seasonal fresh swamps, semi-permanent fresh swamps, teatree swamps and reed
swamps (‘Table 2). Wetlands in these categories contain relatively little open water.

Of the plant taxa recorded, the monocotyledons accounted for nearly 59% of the
total wetland area and dicotyledons about 35% (Appendix). Monocots occurred 1n 97 %
of wetlands and dicots in 89%. The most extensive and commonly recorded monocot
families were Poaceae (23% of wetland area, 82% of wetland number), Cyperaceae
(20% area, 36% number) and Juncaceae (10% area, 75% number). Paspalum distichum
(water couch) dominated the area occupied by the Poaceae and was the most commonly
recorded grass. Eleocharis equisetina was the commonest and by far the most extensive
member of the Cyperaceae. The most extensive and commonly recorded dicot families
were Myrtaceae (17% of wetland area and 36% of wetland number) and Polygonaceae

PROC. LINN. SOC. N.S.W., 111 (3), 1989

148 LOWER CLARENCE WETLANDS

(12% area, 76% number). Melaleuca quinquenervia was the most extensive dicot species,
occupying 17% of the total wetland area. Casuarina glauca was the most commonly
recorded dicot species, occurring in 33% of wetlands, although it is likely that most or all
of the 76% of wetlands containing Persicarza spp. contained FP. hydropiper.

TABLE 2
Extent and decline of wetland types on the far north coast of New South Wales (modified from Goodrick, 1970)#

Original 1969 To
Wetland type area (ha) area (ha) Decline
fresh meadows 22680 ( 2150) 59.4
seasonal fresh swamps ( 7050)
semi-permanent fresh swamps 360 320 Hil. ih
open fresh waters* 450 450 =
teatree swamps 4010 2390 40.4
salt meadows 1380 1380 =
reed swamps 1460 730 50.0
salt flats 530 530 =
mangrove swamps 1860 1780 A)
coastal bogs 570 570 =
coastal Lepironia swamps 320 320 picdue

33620 17670 47.4

# estimates excluded two wetland types: shallow estuarine waters and sheoak swamps.

* open fresh waters is the only floodplain wetland category identified by Goodrick as being dominantly open
water, although relatively small areas of open water occur in other categories on the Clarence floodplain.
Underlined categories are those occurring on the Clarence floodplain.

Cyperaceae and Poaceae were the most diverse families, with 24 and 12 species
respectively, together representing nearly half the total plant taxa. Most plant taxa
occurred in relatively few wetlands and occupied only small percentages of the total wet-
land area (Table 3). The maximum number of native plant taxa recorded in a wetland
was 31, although most wetlands contained relatively few (Table 4).

Four recorded plant species are introduced: Salvinia molesta, Echinochloa crus-gallt,
Nymphaea capensis and Exchhornia crassipes (water hyacinth). At the time of the survey, these
covered about 1.9% of the total wetland area. Water hyacinth was by far the most exten-
sive of these, occupying some 250 ha, and had completely blanketed and infilled some
wetlands.

Several native species, such as Eucalyptus robusta, Juncus kraussi, Sporobolus virginicus
and Trglochin striata, occur only marginally in floodplain wetlands and are more typical
of other wetland types.

The survey provided records of some rare or uncommon plant species. These
include:

— Brasenia schreberr (Gabombaceae): present in 1 wetland; very few records for

coastal New South Wales; listed as 3V by Briggs and Leigh (1988);

— Cyperus odoratus (Cyperaceae): present in 2 wetlands; very few records for the
state;

— Cyperus platystylis (Cyperaceae): present in 1 wetland; a plant of floating organic
mats in northern Australia and a very rare plant in New South Wales (K.
Wilson, Natnl. Herb. N.S.W., pers. comm.); found on a mat of water hyacinth;

— Eleochars philippinensis (Cyperaceae): present in 1 wetland; only one previous
record for the state;

PROC. LINN. SOC. N.S.W., 111 (3), 1989

R. L. PRESSEY 149

TABLES

Frequency of occurrence of plant taxa in wetlands according to Yo number of wetlands and
% total wetland area (bracketed figures indicate number of introduced species in percentage classes)

No. taxa No. taxa
Percentage class (% no. wetlands) (% wetland area)
0-1.0 28(2) 57(
1.1-2.0 9 5(1)
ZF) 5 2
3.1-4.0 1 2
4.1-5.0 8 1
5.1-6.0 Z =
6.1-7.0 2 —
7.1-8.0 2(1) —
8.1-9.0 = —
9.1-10.0 1 1
10.1-20 8 4
21-30 1 —
31-40 2(1) —
41-50 = =
51-60 = =
61-70 = _
71-80 3 ~
81-90 — _
91-100 — =
JUANBILIE, 2¢
Numbers of native plant taxa in wetlands
Taxa No. wetlands % Total no.
> 20 5 0.7
16-20 10 1
11-15 49 6.5
6-10 17 22.6
0-5 520 68.9

755 100.0

— Maunda triglochinoides (Juneaginaceae): present in 34 wetlands over a total area
of 13.5 ha; known range only on the east coast between Wyong and the Brisbane
area (Aston, 1973; Sainty and Jacobs, 1981); considered rare on the New South
Wales central coast by Beadle et al. (1982); generally rarely reported and of
uncertain status (S. Jacobs, Natnl. Herb. N.S.W., pers. comm.).

Despite dry conditions over most of New South Wales in 1982, the results of the
botanical survey should be generally representative of the Clarence floodplain wetlands.
Good rains preceded the February survey when wetland vegetation was well developed
and most basins were full or nearly so. However, some plants would have been missed or
underestimated during the winter part of the survey because of seasonal growth
patterns, absence of inflorescences, or the drier conditions typical of winter in the area.

The majority of wetlands had few trees on their margins (Table 5), a result of the
widespread clearing of the floodplain. Fringing herbaceous emergents were, however,
generally well developed in terms of density, width and the average percentage of
perimeters occupied. The major taxa classed as fringing emergents were Juncus spp. and
Persicaria spp. Others were Carex appressa, Leersia hexandra and Lepironia articulata. Casuarina
glauca and Melaleuca quinquenervia were the most common fringing trees.

PROG. LINN. SOG. N.S.W., 111 (3), 1989

150 LOWER CLARENCE WETLANDS

TABLES
Summary of records for condition of marginal vegetation
A B
Percentage of perimeter with trees Percentage of perimeter with emergent vegetation
Percentage No. wetlands % Votal no. Percentage No. wetlands % Votal no.
91-100 44 5.8 91-100 410 ee
66-90 36 4.8 66-90 V§) 10.5
36-65 58 70 36-65 74 9.8
10-35 65 8.6 10-35 42 5.6
0-9 552 73.1 ()-9 150 19.9
755 100.0 755 100.0

CC. Avcrage density and width of cmergent vegetation

Rating No. wetlands % Votal No.
5 (very dense and wide) 331 43.8
4 69 9.1
3 122 16.2
2 76 10.1
1 (very sparse and narrow) 157 20.8

755 100.0

About 98% of the total number of wetlands were grazed to some extent. Drainage
had directly affected 295 wetlands (39% of total number) and a wetland area of 13,500
ha (92% of total area). Drainage has reduced the persistence and depth of standing
water in these wetlands with consequent changes in the distribution and abundance of
plants and animals. The severity of drainage effects has varied depending on the depth
of drains, the original nature of the wetlands, and the extent of their catchments. The
overall effects of drainage on the Clarence floodplain wetlands are probably underesti-
mated by these statistics for two reasons. Firstly, drainage may have lowered local water
tables and so affected wetlands not directly drained. Secondly, drainage has fragmented
some previously large wetlands into remnant sub-basins that are not directly drained
but have had their water regimes altered.

Few, if any, wetlands on the Clarence floodplain have been completely eliminated
by drainage. The main reason for the inconsistency between this statement and
Goodrick’s (1970) estimates of the decline of fresh meadows and seasonal fresh swamps
in the region (Table 2) is that Goodrick’s survey was primarily concerned with waterfowl
habitat. His estimates of the decline of wetlands therefore included any areas whose
value to waterfowl had been largely eliminated. Many such areas, though, can still be
defined by wetland vegetation.

Wetlands were placed in two broad hydrological categories on the basis of the
nature of their catchments. The first (Category 1) comprises 544 wetlands with an area
of 2456 ha (72% total number, 17% total area) which have small catchments confined to
alluvial flats below the 10 m contour. Their water levels would generally be less stable
and would fall more quickly during dry periods than those in other wetlands.

The second type of wetlands (Category 2) have catchments extending above the 10
m contour beyond the alluvial flats on which they occur. This category includes 211 wet-
lands with a total area of some 12,272 ha (28% of total number, 83% of total area). Most
of these wetlands have catchments smaller than 1000 ha although most of the wetland
area in Category 2 is filled from larger catchments (Table 6). The largest catchment
area, that of Ellis Swamp and Crowsnest Swamp (No. 306 in Fig. 1), 1s 17,200 ha. Most

PROG. LINN. SOG. N.S.W,, 111 (3), 1989

R. L. PRESSEY 151

also have ratios of catchment area/wetland area of less than 100 and nearly 80% of the
wetland area in Category 2 has a ratio of 10 or less (Table 7). Wetlands with large catch-
ments and with large ratios of catchment area/wetland area will generally have more
persistent and stable water levels and will contract more slowly in dry periods. At least
some of these would be important local foci for fauna during droughts and may be
significant in a broader context, particularly for the more mobile species of waterbirds.

TABLE 6
Catchment sizes for wetlands with catchments above the 10 m contour (Category 2)

Catchment No. % Total Wetland % Total
size (ha) wetlands no. Cat. 2 area (ha) area Cat. 2

0-10 62 29.4 112.4 0.9
11-100 66 Sil 3} 421.3 3.4
101-1000 44 20.8 2566.8 20.9
> 1000 39 18.5 - 9171.4 74.7
211 100.0 12272.9 100.0
TABLE 7
Ratios of catchment area/wetland area for wetlands with catchments above the 10 m contour (Category 2)
Catchment/ No. % Votal Wetland % Total
wetland ratio wetlands no. Cat. 2 area area Cat. 2
0-10 106 50.2 9433.0 76.8
11-100 66 31.3 2722.8 22.2
101-1000 26 12.3 95.0 0.8
> 1000 13 6.2 21.1 0.2
211 100.0 122A 100.0

Twenty-one combinations of land uses were recognized in wetland catchments. The
occurrence of each broad land use is summarized in Table 8. Agriculture is by far the
major catchment land use, although areas of natural vegetation are also relatively
common. Seventy-seven percent of catchments are completely under agriculture, and
nearly all of these are restricted entirely to the alluvial flats. Only two catchments are
completely covered by natural vegetation.

TABLE 8
Land use occurrence in wetland catchments
Present (% no. Dominant or sole
Land use catchments) use (% no. catchments)
Agriculture 99.7 5)
Forestry 5.7 Holl
Industrial area 0.5
Mining 357 =
Natural vegetation Zl) UGS)
Urban area 3.0 0.1
Waste disposal 0.5 =

In 95% of wetlands, land tenure is solely or dominantly freehold. One wetland
(Cowans Ponds, No. 257 in Fig. 1) is within a Wildlife Refuge under the National Parks
and Wildlife Act, 1974.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

152 LOWER CLARENCE WETLANDS

The survey reported here is the first comprehensive and detailed description of wet-
lands on the Clarence floodplain. However, the results are accurate for the wetlands and
their catchments only for the period of the field survey and for the dates of the aerial
photography. Subsequent developments and changes in land use will have affected and
will continue to affect the condition of many wetland catchments, the condition of marg-
inal vegetation, and the occurrence of plant taxa and open water. In particular, the
future applicability of the data on plants and open water categories will depend on the
extent of short-term irregular and long-term successional changes in the wetlands. The
nature of these changes ts difficult to predict with the presently poor understanding of
the dynamics of Australian wetlands. Nevertheless, general observations by the author
over several years on the Clarence floodplain suggest that:

— the distribution and abundance of plant taxa in some wetlands alter over
periods of months or years, although the suites of plant species in these areas
generally remain similar;

— the distribution and abundance of plant taxa in other wetlands, particularly
those which are seasonal and have small catchments, appear to change very
little over months or years.

Without research on the vegetation dynamics of these areas, the characteristics of

the wetlands that determine variability or stability of plant distributions, and the nature
and extent of short-term variability, must remain conjectural.

ACKNOWLEDGEMENTS

The study was funded by the N.S.W. National Parks and Wildlife Service as a
consultancy. Phillippa Pressey provided much valuable assistance during compilation
and checking of the survey data and typed the original data base. Chris Ann Urquhart
assisted in the preparation of the tables and drafted the figure. John Porter organized
the analysis of the numerical distribution of open water categories and groups of plant
taxa given in the Appendix. The staff of the National Herbarium of New South Wales,
particularly Surrey Jacobs and Karen Wilson, assisted with the identification of plant
specimens and with advice on their taxonomy and ecology. Surrey Jacobs, Karen
Wilson and an anonymous referee provided helpful comments on the manuscript.

References

ASTON, H. I., 1973. — Aquatic Plants of Australia. Melbourne: Melbourne Univ. Press.

BEADLE, N. C. W., EvANS, O. D. and CAROLIN, R. C., 1982. — Flora of the Sydney Region. 3rd edition. Sydney:
Reed.

BRIGGS, J. D. and LEIGH, J. H., 1988. — Rare or threatened Australian Plants — 1988 Revised Edition. Aust.
Nat. Pks. Wildl. Serv. Spec. Publ. 14.

BROOME, L. S., 1978. — Birds on north coast wetlands. Univ. New England Sch. Nat. Res. Rep. PRY.

CLARENCE RIVER COUNTY COUNCIL, 1978. — Sportsmans Creek — Everlasting Swamp Environmental Impact
Statement. CRCC: Grafton.

——.,, 1980. — Upper Coldstream Area Environmental Impact Statement. CRCC: Grafton.

COASTAL COUNCIL OF NEW SOUTH WALES, 1985. — Coastal Wetlands of New South Wales. Department of
Environment and Planning: Sydney.

GOoDRICK, G. N., 1970. — A survey of wetlands of coastal New South Wales. CSTRO Div. Wildl. Res. Techn.
Memo. 5.

HANNAH, B. C., 1968. — The landscape of the north coast of New South Wales. Univ. New England Dept.
Geogr. Res. Series in Appl. Geogr. No. 14.

Jacoss, S. W. L. and LAPINPURO, L., 1986. — Alterations to the census of New South Wales plants. Zélopea 2:

705-714.
, and PICKARD, J., 1981. — Plants of New South Wales: A Census of the Cycads, Conifers and Angiosperms.
Government Printer: Sydney.

PRESSEY, R. L., 1981. — A Review of Literature on the Floodplain Wetlands of Coastal New South Wales. National

Parks and Wildlife Service: Sydney.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

Reis PRESSEN 153

——, 1987. — A Survey of Wetlands on the Lower Clarence Floodplain, New South Wales. National Parks and Wildlife
Service: Sydney.
, and MIDDLETON, M. J., 1982. — Impacts of flood mitigation works on coastal wetlands in New South
Wales. Wetlands (Australia), 2: 27-44.
SAINTY, G. R. and Jacoss, S. W. L., 1981. — Waterplants of New South Wales. Sydney: Water Resources
Commission.
WEST, R. J., THOROGOOD, C. A., WALFORD, T. R. and WILLIAMS, R. J., 1985. — An estuarine inventory for
New South Wales. N.S.W. Dept. Agric. Fisheries Bull. 2.

PROG. LINN. SOG. N.S.W., 111 (3), 1989

LOWER CLARENCE WETLANDS

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PROC. LINN. SOC. N.S.W., 111 (3), 1989

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Wetlands of the Lower Macleay Floodplain,
Northern Coastal New South Wales

Reo Ie JERUSS SINC
(Communicated by P. ADAM)

PRESSEY, R. L. Wetlands of the lower Macleay floodplain, northern coastal New South
Wales. Proc. Linn. Soc. N.S.W. 111 (3), 1989: 157-168.

A survey of wetlands on the lower Macleay floodplain in 1984 involved mapping
and description of 432 wetlands with a total area of nearly 12,800 ha. Most wetlands
were less than 10 ha and the three largest accounted for 73% of the total area. One
hundred and eighteen plant taxa and two categories of open water were recorded. Open
water occupied only 2.5% of the total wetland area but some open water occurred in
54% of wetlands. The most extensive and frequently occurring plant families were
Polygonaceae (covering 26% of the wetland area, occurring in 97% of the total number
of wetlands), Poaceae (28% area, 91% number), Juncaceae (10% area, 91% number),
Cyperaceae (20% area, 63% number) and Myrtaceae (6% area, 21% number). Broad
fringing bands of herbaceous plants were typical of the wetlands. Most wetlands had
few trees on their margins due to clearing. About 99% of wetlands were grazed.
Drainage affected 51% of the total number and 96% of the total wetland area. Most
wetlands are seasonal and have relatively small catchments restricted to the alluvial
flats. Twenty-one percent of wetlands have catchments on adjacent hillslopes and
ranges. Agriculture was by far the dominant land use in wetland catchments. Most wet-
lands were freehold and only one is covered (partly) by a reserve for nature
conservation.

R. L. Pressey, New South Wales National Parks and Wildlife Service, PO. Box 1967, Hurstville,
Australia 2220; manuscript received I November 1988, accepted for publication 15 March 1989.

KEY WORDS: Wetlands, survey, plant taxa, catchments, Macleay floodplain.

INTRODUCTION

The wetlands of the Macleay floodplain have not previously been mapped or
surveyed in a detailed and comprehensive way. In a broad-scale survey of the State’s
coastal wetlands, Goodrick (1970) mapped and classified many on the Macleay flood-
plain, including all the major ones. His study outlined the types and distribution of
coastal wetlands and estimated their decline up to 1969. However, it lacked the detailed
information needed to decide priorities for conservation and management. The only
other inventory of wetlands on the Macleay floodplain was by the Coastal Council of
New South Wales (1985) which mapped many of them for zoning purposes without
describing or classifying them. Another broad survey by West ef al., (1985) dealt
exclusively with the estuarine wetlands of the Macleay and other parts of the coast.

There are few detailed studies on the flora and fauna of the Macleay floodplain wet-
lands and none of these has covered more than a few of the total number of wetlands.
Broome (1978) identified the habitat associations of waterbirds and studied the
dynamics of their populations at a few selected sites. The remaining studies on the wet-
lands of the floodplain are mainly concerned with geomorphology and soils (Pressey,
1981).

This paper summarizes the results of a detailed inventory and description of all the
wetlands on the lower Macleay floodplain (see Pressey, 1987). The survey was part of a
series of studies on coastal wetlands undertaken for the New South Wales National Parks
and Wildlife Service. The area and aims of the surveys and the types of data collected
are outlined in the accompanying paper on the Clarence floodplain wetlands (Pressey,

1989).

PROC. LINN. SOC. N.S.W., 111 (3), 1989

158 LOWER MACLEAY WETLANDS

STUDY AREA

The Macleay River floodplain is situated on the mid-north coast of New South
Wales, approximately 350 km north-east of Sydney (Fig. 1). As for the Clarence survey,
the lower Macleay floodplain was defined as alluvium below the 10 m contour. This
boundary contains virtually all of the alluvial wetland in the Macleay system and en-
closes the extensive deltaic alluvial plain that has formed during and since the last rise in
sea level. The floodplain above the 10 m contour 1s restricted to a narrow strip along the
river and the wetlands are generally small and scattered. The survey was also restricted
to predominantly fresh wetlands and excluded estuarine and most brackish areas.

SOUTH WEST ROCKS

<— STUDY AREA

Frederickton

Y,
5° BY
EY Gf &

a
2

KEMPS
7

Sherwood 263555
Aldavilla § bey 4
I yz —

Fig. 1. The study area. Numbers are those given to named wetlands during the survey. The names corres-
ponding to these numbers are: 10 Christmas Creek; 69 Doughboy Swamp/Sevenoaks Swamp/Clybucca
Swamp; 133 Cooroobongatti Swamp; 261 Chapmans Creek; 263 Chapmans Creek; 265 Chapmans Creek;
275 East Kempsey Swamps/Bridge Creek; 278 Pola Creek; 350 Belmore Swamp; 353 Swan Pool/Kinchela
Swamp.

The study area occupies about 400 sq. km (Fig. 1). To the north of the river it
extends upstream as far as Aldavilla. On this side, a large area of alluvium lies adjacent
to Christmas Creek, immediately north of Kempsey, but the bulk of the northern flood-
plain is to the north and east of Frederickton where continuous alluvium extends for
some 140 sq. km. The dominant wetland area here is the large Doughboy, Sevenoaks or

PROC. LINN. SOC. N.S.W., 111 (3), 1989

R. L. PRESSEY 159

Clybucca Swamp in the north-west of the study area. ‘To the south of the river the study
area extends upstream almost to Sherwood. Most of the southern alluvium lies in a con-
tinuous tract of about 180 sq. km downstream of South Kempsey. The largest wetland
areas here are the Swan Pool, formerly known as Kinchela Swamp, and Belmore
Swamp.

The main land use on the Macleay floodplain is grazing for dairying and beef.
Relatively small areas have been planted to poplars and to crops such as sorghum and
corn. As on the Clarence floodplain, drainage of wetlands began late in the nineteenth
century to improve pasture production. Drainage trusts or unions were formed around
the turn of the century to co-ordinate drainage schemes. From the late 1950s, under
flood mitigation schemes, drains, floodgates and levees were constructed and integrated
to minimize damage to urban and agricultural land from flooding. They extended and
intensified the alteration of wetlands caused by earlier drainage.

Goodrick (1970) found that about 49% of all wetlands on the mid-north coast,
which includes the Macleay floodplain, had been lost or significantly altered before
1970, mainly to drainage and flood mitigation. This decline or alteration includes about
14,800 ha of shallow floodplain wetlands, representing about 68% of the wetlands lost or
altered. The impacts on floodplain wetlands are likely to be more extensive today since
flood mitigation on the lower Macleay has continued since Goodrick’s survey (Pressey
and Middleton, 1982).

METHODS

Initial identification of wetlands was from black and white or colour aerial photo-
graphs taken between 1979 and 1981 at scales of 1:25 000 and 1:50 000. The wetlands
were each visited at least once during two field surveys (6.12.83 — 21.12.83 and 6.1.84 —
27.1.84).

Of thirteen attributes of each wetland recorded in the field and/or remotely from
maps and aerial photographs, six were used only for ranking the wetlands for conser-
vation (see Pressey, 1987). The remaining seven — size, vegetation/habitats, marginal
vegetation, alteration, catchment areas, catchment land use, and tenure — were used
descriptively and are discussed in this paper.

Methods of recording the seven descriptive attributes are given in the preceding
paper on the Clarence wetlands (Pressey, 1989). Differences in recording plant taxa in
the Macleay wetlands were that more time was spent searching for plants, species in the
families Azollaceae and Lemnaceae were included, and all plants were identified at least
to species level. Authorities for plant names are those in Jacobs and Pickard (1981) as
amended by Jacobs and Lapinpuro (1986), unless indicated otherwise.

RESULTS AND DISCUSSION

The survey covered 432 wetlands with a total area of 12,772 ha. Most of the wet-
lands are small, 90% being 10 ha or smaller (Table 1). These small wetlands account for
less than 6% of the total wetland area. The bulk of the wetland area (89%) is made up by
the relatively small number (3% of the total) greater than 100 ha in size. The three
largest wetlands account for 73% of the total wetland area.

The recorded plant taxa, including a hybrid Juncus, an undescribed species of
Myrnophyllum, and two forms of Persicaria strigosa, totalled 118. Their occurrence in the
study area, with that of the two recorded categories of open water, is summarized in the
Appendix.

The two forms of Ferszcarna strigosa differed in features and habitat. P. strigosa FORM
1 was the less common. It is characterized by large green leaves with cordate bases,

PROC. LINN. SOC. N.S.W., 111 (3), 1989

160 LOWER MACLEAY WETLANDS

bristles beneath the dense glandular hairs on the peduncles, and white flowers in a large,
compact, ovate spike. It was found generally along the edges of relatively permanent,
steep-sided channels and in other sheltered, damp areas that are rarely inundated. P
strigosa FORM 2 has reddish leaves that are more lanceolate than FORM 1, frequently
with hastate bases. In addition, there are no bristles beneath the glandular hairs on the
peduncles and the spike of white flowers is linear to ovate and occasionally interrupted.
It was found growing in dense meadows in relatively persistent standing water, particu-
larly in wetlands with large catchments or in those adjacent to perennial streams.
FORM 2 is the more similar to P sp. B in terms of leaf shape and inflorescence
structure.

TABLE 1
Size distribution of wetlands
No. of % Total Wetland % Total
Area (ha) wetlands no. area (ha) area
0-1.0 190 44.0 107.5 0.8
{1-5-0 162 378 368.8 2.9)
5.1-10.0 37 8.6 254.5 2.0
10.1-50 28 6.5 526 4.1
51-100 3 0.7 200 1.6
101-500 8 1.8 1415 iil, il
501-1000 1 0.2 530 41
> 1000 3 0.7 9370 73.4
432 100.0 12771.8 100.0

Open water was relatively unimportant in terms of the overall wetland area
occupied (Appendix). Some shallow open water occurred in about half the number of
wetlands but totalled, at the time of the survey, only about 290 ha or 2.3% of total wet-
land area. Deep open water was much more restricted in distribution and extent. The
scarcity of open water in the wetlands of the mid-north coast generally is indicated by
estimates from Goodrick’s (1970) report (Table 2). The five categories of Goodrick’s
classification that make up the bulk of the wetland area in the region, and on the
Macleay floodplain, contain relatively little open water and are dominated by her-
baceous emergents and by trees.

Completion of all the field work in mid-summer following heavy rains provided
ideal conditions for the identification of a large number of wetland plant taxa. Some
shallow seasonal wetlands contained so much water that the results of the survey may be
atypical to some extent. Of the plant taxa recorded, one 1s a liverwort, seven are ferns, 67
are monocotyledons and 43 are dicotyledons. The most diverse families were Cyper-
aceae (30 taxa), Polygonaceae (12), Juncaceae (11) and Poaceae (10). The largest number
of native taxa recorded in a wetland was 46, although most wetlands contained relatively
few (Table 3). There is a significant relationship between wetland size and number of
plant taxa in the wetlands on the Macleay floodplain (Margules et al., 1988) although
there is considerable variability in the number of taxa in wetlands of any size. Most taxa
occurred in relatively few wetlands and occupied a relatively small overall area (Table 4).

Monocotyledonous plants occupied some 62% of the total wetland area and
occurred in all but one wetland (Appendix). The most important monocot families, in
terms of extent and frequency of occurrence, were Poaceae (28% of total wetland area,
91% of total number of wetlands), Juncaceae (10% area, 91% number) and Cyperaceae
(22% area, 63% number). Paspalum distichum (water couch) and Juncus polyanthemus x
usitatus were the most extensive and frequently occurring taxa in the Poaceae and

PROG. LINN. SOG. N.S.W., 111 (3), 1989

R. L. PRESSEY 161

_Juncaceae, respectively. Eleocharis equisetina made up most of the area occupied by the
Cyperaceae and was one of the most frequently occurring sedges. Dicotyledons
occupied 35% of the wetland area and occurred in all but three wetlands. Polygonaceae
(26% area, 97% number) and Myrtaceae (6% area, 21% number) were the major dicot
families. Persicarta hydropiper and Melaleuca quinquenervia were, respectively, the most
extensive and commonly recorded species in these families.

ARIE
Extent and decline of wetland types on the mid-north coast of New South Wales (modified from Goodrick, 1970)#
Original 1969
Wetland type area (ha) area (ha) % Decline
fresh meadows 18590 ( 2430) T9a0
seasonal fresh swamps ( 1340)
semi-permanent fresh swamps 120 120 —
open fresh waters* 40 40 -
shallow saline lagoons 650 650 —
teatree swamps 7490 6560 12.4
salt meadows 3520 3520 —
reed swamps 2880 810 71.9)
salt flats 160 160
mangrove swamps 1170 1170 =
coastal bogs 1940 1940 -
coastal Lepironia swamps 40 40 =
36600 18780 48.7

# estimates excluded two wetland types: shallow estuarine waters and sheoak swamps.

* open fresh waters is the only floodplain wetland category identified by Goodrick as being dominantly open
water, although relatively small areas of open water occur in other categories on the Macleay floodplain.
Underlined categories are those occurring on the Macleay floodplain.

TABLE 3
Numbers of native plant taxa in wetlands
Taxa No. wetlands % Total no.
> 30 4 0.9
26-30 3 0.7
21-25 9 Dall
16-20 31 12
11-15 101 23 Ea:
6-10 174 40.2
0-5 110 25.5
432 100.0

Twenty of the species recorded are introduced. They occupied about 180 ha or
1.5% of total wetland area at the time of the survey. The most extensive were Ezchhornia
crassipes (94 ha), Salvinia molesta (33 ha) and Echinochloa crus-galli (31 ha). The first two
mentioned are particularly troublesome floating weeds and had completely blanketed
and infilled some wetlands. Cyperus brevifolius may be native to parts of northern Aus-
tralia although it is considered to be introduced to the study area (K. Wilson, Natnl.
Herb. N.S.W., pers. comm.).

Four recorded species, Commelina cyanea, Cyperus polystachyos, Pseudognaphalium luteo-
album and Ranunculus plebeius, are widely distributed in non-wetland habitats, particu-

PROC. LINN. SOC. N.S.W., 111 (3), 1989

162

LOWER MACLEAY WETLANDS

larly in higher rainfall areas. A further six species, Bacopa monniert, Bolboschoenus cald-
welli, Juncus kraussu, Schoenoplectus litoralis, Sporobolus virginicus and Triglochin striata, are
typical of brackish and estuarine habitats and occur only marginally in freshwater flood-
plain wetlands.

TABLE 4

Frequency of occurrence of plant taxa in wetlands according to Yo number of wetlands and
% total wetland area (bracketed figures indicate number of introduced species in percentage classes)

No. taxa No. taxa
Percentage class (% no. wetlands) (% wetland area)
0-1.0 39(9) 107(20)
1.1-=2,,0 14(1) 1
2163-0 12(1) 3
371-420 4(2) =
4.1-5.0 4 1
5.1-6.0 (3) 1
6.1-7.0 5 =
7.1-8.0 3) =
8.1-9.0 4 2
9.1-10.0 2 2
10.1-20 12(2) 1
21-30 5(1) =
31-40 1 =
41-50 2(1) =
51-60 1 —
61-70 2 =
71-80 1 =
81-90 2 =
91-100 = —

Four rare or uncommon plant species were recorded during the field survey:

Cyperus platystylis (Cyperaceae): present in six wetlands over a total area of 0.1
ha; very rare in New South Wales although more common in northern Aus-
tralia where it occurs on floating organic mats (K. Wilson, Natnl. Herb.
N.S.W., pers. comm.); recorded only on mats of the introduced Salvinia molesta
and Exchhornia crassipes;

Eleocharis philippinensis (Cyperaceae): present in one wetland; third and
southernmost record for the State;

Maundia triglochinoides (Juncaginaceae): present in 30 wetlands and over a total
area of 12.1 ha; known range only between Wyong and the Brisbane area
(Aston, 1973; Sainty and Jacobs, 1981); considered rare on the New South
Wales central coast by Beadle et al. (1982); generally rarely collected and of
uncertain status (S. Jacobs, Natnl. Herb. N.S.W., pers. comm.);

Potamogeton javanicus (Potamogetonaceae): present in one wetland; only known
from the central and north coast regions of the State (Sainty and Jacobs, 1981)
and recorded very rarely over this range although it 1s widespread in northern
Australia and overseas (Aston, 1973); given as rare on the central coast by
Beadle et al., (1982) and rarely collected from the Sydney region in recent years
(R. Coveny, Natnl. Herb. N.S.W., pers. comm.).

The majority of wetlands had dense, broad bands of herbaceous emergents around
most of their perimeters (Table 5). The major fringing emergents, by far, were Persicarza
hydropiper and Juncus polyanthemus x usitatus. Few wetlands had more than 10% of their
margins lined with trees, the result of clearing for grazing. The commonest fringing
trees were Casuarina glauca and Melaleuca quinquenervia.

PROG. LINN. SOC. N.S.W., 111 (3), 1989

R. L. PRESSEY 163

TABLE 5
Summary of records for condition of marginal vegetation
A B
Percentage of perimeter with trees Percentage of perimeter with emergent vegetation
Percentage No. wetlands % ‘Yotal no. Percentage No. wetlands % ‘Votal no.
91-100 1() Dos 91-100 at 75.0
66-90 12 2.8 66-90 19 4 at
36-65 15 32) 36-65 32 7.4
11-35 27 6.2 11-35 12 2.8
0-10 368 85.2 0-10 45 10.4
432 100.0 432 100.0

C. Average density and width of emergent vegetation

Rating No. wetlands % Yotal No.

5 (very dense and wide) 316 T32

4 26 6.0

3 26 6.0)

2 19 4.4

1 (very sparse and narrow) 45 10.4
432 100.0

About 99% of the wetlands were grazed to some extent by cattle. At least five wet-
lands had been separated from tidal influence by floodgates or earthen block banks.
Drainage had directly affected 220 wetlands (51% of total number) and a wetland area
of 12,236 ha (96% of total area). Additional, indirect effects of drainage on other wet-
lands include separation into undrained sub-basins and could include local lowering of
water tables.

The actual hydrological effects of drainage would vary with factors such as the
depth of drainage and of water tables and the size of catchments. In general, however,
drained wetlands will have reduced storage capacity and will dry more readily com-
pared to undrained ones. It 1s unlikely that any wetlands on the lower Macleay flood-
plain have been completely eliminated by drainage, despite Goodrick’s (1970) estimates
of wetland decline in the region (Table 2). His estimates are based on the elimination of
value to waterfowl whereas wetlands were defined in this study by the occurrence of
plants adapted to at least seasonal inundation. Many wetlands defined in this way are no
longer of value as waterfowl habitat. Some 33% of the wetland area covered by the
survey reported here was occupied by two taxa (Juncus polyanthemus x usitatus and
Persicaria hydropiper) typical of marginal wetland conditions. Wetland drainage has
probably been a major cause of the extensive distribution of these taxa.

Two broad hydrological categories of wetlands were distinguished from the nature
of their catchments. The first (Category 1) contains 342 wetlands (79% of total number)
with a combined area of nearly 4800 ha (37% of total area) which have catchments
extending no higher than the 10 m contour. They are filled only by groundwater and
localized runoff from the surrounding alluvial flats and will generally have less stable
water levels and will dry more readily than those in Category 2. The second type of wet-
lands (Category 2), with additional drainages on the bedrock slopes surrounding the
floodplain, number 90 (21% of total number) and have an overall area of about 8,000 ha
(63% of total area).

Most wetlands in Category 2 have catchments on bedrock smaller than 100 ha but
the few large wetlands which make up most of the area of this type are fed by catchments

PROG. LINN. SOG. N.S.W., 111 (3), 1989

164 LOWER MACLEAY WETLANDS

larger than 1000 ha (Table 6). The catchment sizes for these wetlands are 1100 ha (East
Kempsey Swamps, No. 275 in Fig. 1), 1790 ha (Belmore Swamp, No. 350), 5060 ha
(Christmas Creek, No. 10) and 13,400 ha (Doughboy Swamp, No. 69). The other very
large catchment covers 1880 ha and supplies wetland No. 249 which has an area of only
about 1 ha. About 90% of wetlands in Category 2 have catchment area/wetland area
ratios of 100 or less (Table 7). About 95% of the area in this type of wetland is fed by
catchments that are at most 10 times larger than the individual wetlands. Larger catch-
ments and larger ratios of catchment area/wetland area will generally confer greater
permanence on wetlands or on particular parts of large, complex basins. Water levels in
these wetlands would generally be more persistent because of the greater effectiveness of
any rainfall in refilling the basins and perhaps because of greater base flow from ground-
water in large catchments.

TABLE 6
Catchment sizes for wetlands with catchments above the 10 m contour (Category 2)

Catchment No. % Total Wetland % Total

size (ha) wetlands no. Cat. 2 area (ha) area Cat. 2
0-10 43 47.8 67.9 0.9
11-100 29 32.2 78). 0.9
101-1000 13 14.4 503.0 6.3
> 1000 5 5.6 7340.8 Oil
90 100.0 7986.8 100.0
TABLE 7
Ratios of catchment area/wetland area for wetlands with catchments above the 10 m contour (Category 2)
Catchment/ No. % Total Wetland % Total
wetland ratio wetlands no. Cat. 2 area area Cat. 2
0-10 54 60.0 7553.8 94.6
11-100 28 31.1 ANT fl 52)
101-1000 7 7.8 14.5 0.2
> 1000 1 1.1 0.8 =
90 100.0 7986.8 100.0

Table 8 summarizes the occurrence of seven broad land uses in the catchments of
the Macleay floodplain wetlands. Every wetland catchment contains some agricultural
use and agriculture is the dominant or sole use in 93% of catchments. Areas of natural
vegetation occur in 21% of catchments but occupy the largest parts of only 4% of catch-
ments. None of the wetlands surveyed had completely natural catchments. Forestry,
industrial areas, mining (only surface extraction of sand, gravel etc. in the case of the
Macleay survey area), urban areas and waste disposal occurred in relatively few catch-
ments and rarely or never occupied the largest proportions.

The tenure of about 98% of the wetlands was solely or dominantly freehold. A
small part of the Swan Pool (wetland No. 353) is within Hat Head National Park.

The results reported here are estimates that are strictly valid only for the period of
the field survey and for the dates of the aerial photography. The condition of wetlands
and their catchments and the tenure of the wetlands will have since changed to some
extent with new developments and changes in land use. In particular, short-term
dynamics of wetland vegetation will have altered the occurrence of plant species and
open water. Further surveys of the wetlands and research on the temporal variability of
coastal freshwater wetlands are necessary to quantify and explain such changes.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

R. L. PRESSEY 165

TABLE 8
Land use occurrence in wetland catchments
Present (% no. Dominant or sole
Land use catchments) use (% no. catchments)
Agriculture 100.0 93.3
Forestry 1.4 0.9
Industrial area 0.7 —
Mining 2
Natural vegetation 2AES 4.2
Urban area 5.8 1.6
Waste disposal 1.4
ACKNOWLEDGEMENTS

The study was funded by the N.S.W. National Parks and Wildlife Service as a
consultancy. Phillippa Pressey provided much valuable assistance in the compilation
and checking of the survey data and typed the original data base. Chris Ann Urquhart
assisted in the preparation of the tables and drafted the figure. John Porter organized
the analysis of the numerical distribution of open water categories and groups of plant
taxa given in the Appendix. Staff of the National Herbarium of New South Wales are
thanked for processing and identifying plant specimens. Thanks are due to Karen
Wilson, in particular, for advice on identification. Karen Wilson, Surrey Jacobs and an
anonymous referee provided helpful comments on the manuscript.

References

ASTON, H.I., 1973. — Aquatic Plants of Australia. Melbourne: Melbourne Univ. Press.
BEADLE, N. C. W., Evans, O. D. and CAROLIN, R. C., 1982. — Flora of the Sydney Region. 3rd Edition. Sydney:

Reed.
BROOME, L. S., 1978. — Birds on north coast wetlands. Univ. New England Sch. Nat. Res. Rep. PR9.
COASTAL COUNCIL OF NEW SOUTH WALES, 1985. — Coastal Wetlands of New South Wales. Department of

Environment and Planning: Sydney.
GooprRICck, G. N., 1970. — A survey of wetlands of coastal New South Wales. CSTRO Div. Wildl. Res. Techn.
Memo. 5.
Jacoss, S. W. L. and LaPINPpuRO, L., 1986. — Alterations to the census of New South Wales plants. Télopea 2:
705-714.
, and PICKARD, J., 1981. — Plants of New South Wales: A Census of the Cycads, Conifers and Angiosperms.
Government Printer: Sydney.
MARGULES, C. R., NICHOLLS, A. O. and PREsSSEY, R. L., 1988. — Selecting networks of reserves to
maximise biological diversity. Bzol. Cons., 43: 1-13.
PRESSEY, R. L., 1981. — A Review of Literature on the Floodplain Wetlands of Coastal New South Wales. National
Parks and Wildlife Service: Sydney.
——., 1987. — A Survey of Wetlands of the Lower Macleay Floodplain, New South Wales. National Parks and Wildlife
Service: Sydney.
——, 1989. — Wetlands of the lower Clarence floodplain, northern coastal New South Wales. Proc. Linn. Soc.
N.S.W,, 111: 143-155.
, and MIDDLETON, M. J., 1982. — Impacts of flood mitigation works on coastal wetlands in New South
Wales. Wetlands (Australia), 2: 27-44.
SAINTy, G. R. and Jacoss, S. W. L., 1981 — Waterplants of New South Wales. Sydney: Water Resources
Commission.
WEsT, R. J., THOROGOOD, C. A., WALFORD, T. R. and WILLIAMS, R. J., 1985 — An estuarine inventory for
New South Wales. N.S.W. Dept. Agric. Fisheries Bull. 2.

PROC. LINN. SOG. N.S.W,, 111 (3), 1989

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PROC. LINN. SOC. N.SW., 111 (3), 1989

The Ordovician-Silurian Stratigraphy of the
Cudgegong-Mudgee District, New South Wales

JOHN W. PEMBERTON

PEMBERTON, J. W. The Ordovician-Silurian stratigraphy of the Cudgegong-Mudgee
district, New South Wales. Proc. Linn. Soc. N.S.W. 111 (3), 1989: 169-200.

Ordovician and Silurian rocks of the Cudgegong-Mudgee district crop out in a
narrow belt separating Devonian sequences on the northern Capertee High.

The basal unit consists of Late Ordovician basaltic and andesitic lava and their
fragmental equivalents (Cudgegong Volcanics) which are unconformably overlain by a
thick Wenlockian to Ludlovian sequence of shallow marine to emergent units. The
lowermost Willow Glen Formation (fossiliferous clastic sediments and limestone) was
deposited in a coastal environment affected by a series of transgressive/regressive cycles.
It is overlain by a thick and persistent sequence of dacite lava and breccia (Windamere
Volcanics) with associated mass flow detritus (Zoolamanang Formation), followed by
volcanic quiescence with continuing shallow marine sedimentation (Millsville
Formation).

The Ordovician-Silurian rocks occupy the core of a shallowly plunging northwest-
trending anticline, the northeast limb of which is overturned. The rocks of this belt are
disconformably overlain by Early Devonian units, a continuation of shallow marine
conditions on the northern Capertee High, whereas the fluviatile Late Devonian
Lambie Group overlies the strata with slight angular unconformity.

J. W. Pemberton, Department of Geology, University of Wollongong, PO. Box 1144, Wollongong,
Australia. 2500; manuscript received 12 July 1988, accepted for publication 19 April 1989.

LOCATION

Mudgee is 260km northwest of Sydney in the central tablelands of New South
Wales (Fig. 1). The village of Cudgegong, 35km to the southeast, and surrounding
properties have been resumed by the Water Resources Commission as part of the
Windamere Dam and Recreation Park, along the Cudgegong River valley. Con-
sequently, parts of the study area including Cudgegong township have, since 1984, been
flooded by the infilling Windamere Dam.

GEOLOGICAL SETTING

The Cudgegong-Mudgee district lies near the northeast margin of the Lachlan
Fold Belt and comprises sequences mainly deposited on the Capertee High (Fig. 1). In
this paper, a detailed stratigraphy is presented for the southeast-trending belt of Late
Ordovician to Late Silurian rocks which crops out along the Cudgegong River valley
from Mudgee to Cudgegong. The belt is flanked to the northeast and southwest by
Devonian sequences whereas in the south, the belt is intruded by the Carboniferous
Aarons Pass Granite and unconformably overlain by Permian strata.

There are no known Cambrian rocks in the northeast Lachlan Fold Belt. However,
by the Early Ordovician deep water flysch sequences were common with scattered mafic
volcanic centres and associated shoalwater limestones developing in the Parkes, Welling-
ton, and Molong areas (Cas, 1983). These conditions continued into the Late Ordo-
vician where the dominant feature in the eastern Lachlan Fold Belt was the
north-northwest-trending Molong High — a volcanic are producing andesitic centres
with flanking shoalwater limestones and volcaniclastics in an otherwise flysch-
dominated environment (Powell, 1983b; Cas, 1983).

Deformation at the end of the Ordovician apparently continued into the early
Middle Silurian (Crook et al., 1973) and heralded the onset of a tensional regime in the

PROC. LINN. SOC. N.S.W., 111 (3), 1989

170 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

post Devonian igneous
and sedimentary rocks
[- | Devonian sedimentary rocks

= MILLSVILLE FORMATION

ve TOOLAMANANG FORMATION

rhyolite

w/e WINDAMERE VOLCANICS
= dacite

SILURIAN
fac WILLOW GLEN FORMATION

To Sydney

ORDOVICIAN fa] CUDGEGONG VOLCANICS

Fig. 1. Ordovician-Silurian geology of the Cudgegong-Mud¢gee district.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 171

Late Silurian which produced regionally extensive silicic magmatism and extensional
marine basins (Pickett, 1982b; Powell, 1983b). In the northeast, the deformation
initiated rifting of the Molong High producing the opening of the Hill End Trough as a
deep-water basin flanked to the east by a rifted fragment which produced the Late
Ordovician basement for the developing Capertee High (Gilligan and Scheibner, 1978).
Limestones, epiclastics, and extensive silicic volcanics represent Late Silurian shallow
marine deposition on both the Molong and Capertee Highs whereas detritus for the
infilling Hill End Trough was derived from the volcanism (Pickett, 1982b). On the east
margin of the Lachlan Fold Belt, this configuration continued through the Early
Devonian.

By the Middle Devonian, much of the Lachlan Fold Belt had been uplifted and
intensely deformed, and proven Middle Devonian strata are limited to the Capertee
High (Pickett, 1982b). However, there is a marked facies difference between the Early
and Late Devonian sequences, accompanied by low angle discordant contacts along the
northeast margin. The Late Devonian fluviatile and marine conditions (Powell, 1983a)
continued into the Early Carboniferous and the orogenic history of the Lachlan Fold
Belt concludes with terminal deformation in the late Early Carboniferous (Powell and
Edgecombe, 1978; Cas, 1983).

PREVIOUS GEOLOGICAL INVESTIGATIONS

The first major treatment of the geology of the district was that of Game (1935) in
which he mapped a wide tract of Silurian and Devonian strata from Mudgee to Aarons
Pass and Kandos. He considered that a central belt of Upper Devonian rocks was
faulted against Upper Silurian sequences. Mapping by the present author in the south-
west Silurian belt has shown that: the sequence includes both Ordovician and Late
Silurian rocks; conformable and unconformable contacts exist; the Ordovician-Silurian
rocks are folded into a large scale anticline; and the strata may be locally overturned.
Clearly this revised stratigraphy greatly affects part of Game’s interpretation.

Wright (1966) produced a major contribution to the understanding of the Devonian
stratigraphy and faunas and, in particular, to the subdivision of Game’s Upper
Devonian sequences on the basis of Early and Late Devonian faunas. In addition, he
recognized that many of Game’s limestones in the Queens Pinch area were, in fact,
Early Devonian.

On the Dubbo 1:250 000 geological sheet, Offenberg e¢ al. (1971) portrayed, in the
Cudgegong-Mudgee district, the Devonian sequences of Wright (1966) flanking a
central area in which they recognized the Ordovician Sofala Volcanics and the Siluro-
Devonian Gulgamree Beds. However, mapping by Powis (1975), Michie (1975) and
Pemberton (1977) raised doubts as to the validity of this nomenclature. Recent mapping
by the author indicates that the proposed extent of the Sofala Volcanics of Offenberg et
al. includes both Ordovician basaltic sequences and Silurian dacitic sequences. In
addition, Wright (pers. comm.) considers their use of the Gulgamree Beds to far exceed
his original description of the unit which, in any case, is probably a lateral equivalent of
the Mullamuddy Formation in the Queens Pinch Belt. Most of the sedimentary rocks
placed in the Gulgamree Beds by Offenberg et al. (1971) are now mapped as the Willow
Glen Formation (Pemberton, 1980b).

CUDGEGONG VOLCANICS (Lavers 1960)

The Cudgegong Volcanics, the oldest rocks in the district (Fig. 2), were first
mentioned by Lavers (1960) and subsequently by McManus ¢t al. (1965). These rocks, to
the southeast of Cudgegong, have subsequently been described by Pemberton (1980a).

PROC. LINN. SOC. N.S.W., 111 (3), 1989

172 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

Further mapping has revealed four separate outcrop areas of the Cudgegong Volcanics
(Lawes, 1 IE, WE ULE ual NY iheoyan Jes, 3),

PERMIAN SHOALHAVEN

CARBONIFEROUS AARONS PASS GRANITE

MT FROME TO KANDOS
DEVONIAN etal
QUEENS PINCH BELT

Ludlovian
WINDAMERE TOOLAMANANG

SILURIAN
VOLCANICS FORMATION

Wenlockian
WILLOW GLEN FORMATION

ORDOVICIAN CUDGEGONG VOLCANICS

Fig. 2. Stratigraphy of the Cudgegong-Mudgee district.

Stratigraphic Relationships

In each of the four outcrop areas, the Cudgegong Volcanics are unconformably
overlain (the contact is not exposed) by the Willow Glen Formation, or are faulted
against younger rocks. The base of the Volcanics is not exposed.

The areas consist mainly of fine- to medium-grained andesitic arenite, basaltic
rocks being generally rare. Exceptions include: the southeast area (IV) where abundant
basaltic textures and compositions are modified by the contact metamorphism of the
Aarons Pass Granite; common basaltic arenite throughout the southwest area (II); and
the larger central area (III, Fig. 3) which provides the best exposures of the basaltic
rocks and andesitic lava. In the northwest of the latter area, lava, with probable pillows,
fine- to coarse-grained arenite and breccia together with andesite lava and possible

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 173

syenite sills suggest proximity to a basaltic source (Cas and Wright, 1987). To the
southeast, near Limestone Creek, there is a fining in fragment sizes within near con-
tinuous basaltic exposures and here the fine- to medium-grained basaltic arenite may
indicate a more distal aspect from the possible northwest source. The basaltic exposure,
near the southwest margin (GR 615695), consists of lava, with probable pillows, arenite
and minor breccia.

To Mudgee Outcrop areas | to IV for

_the Cudgegong Volcanics

symbols as
for Fig. 1

A——B Representative
cross-section
new Cudgegong-—
Mudgee road
old Cudgegong-
Mudgee road

oD
7a

<
cS
Re)
WB Go

8c,

Toolamanang Formation

Windamere Volcanics

Willow Glen Formation

basaltic rocks
syenite

basaltic breccia
with limestone clasts

shale

andesitic lava

andesitic arenite

Fig. 3. Geology of the central area (III) of the Cudgegong Volcanics.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

174 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

All areas display rapid lateral lithological variation and recognition of marker
horizons is difficult. The exception is in the northwest of the central area (Fig. 3) where a
number of distinctive horizons of the less common rock types are exposed. These

include andesite lava, shale, breccia with abundant limestone clasts and syenite
horizons.

TABLE 1
Petrography of the main rock types in the Cudgegong Volcanics

Porphyritic with phenocrysts of subhedral to euhedral clinopyroxene
(glomeroporphyritic, twinned, zoned, diopside to augite, 11 to 34%) to 7mm
across; subhedral to euhedral plagioclase (albite, 0 to 18%) to Smm long;

and rare subhedral amphibole (pargasite, X=colourless, Y=light green,
Z=green/brown) to 2mm across. Groundmass (50 to 59%) pilotaxitic to
cryptocrystalline (usually recrystallised) of albite laths, clinopyroxene and

rare amphibole subhedra, tremolite, sphene, chlorite, calcite, epidote and
pyrite. Rare ovoid amygdales contain chlorite and calcite. Clinopyroxene
chemistry and significance detailed in Pemberton and Offler (1985); relatively
unaltered with chlorite and calcite along fractures in cores, and tremolite rims
On more altered grains. Albite strongly altered to sericite, chlorite and
calcite. Initial glassy groundmass devitrified and now dominated by chlorite and
sphene. Calcite, chlorite, epidote, albite, prehnite, pumpellyite and quartz
common in porphyroblastic aggregates.

basaltic Very fine— to very coarse-grained (0.01 to 2mm), moderately sorted in ash

arenite fractions to poorly sorted in coarser rocks, immature. Angular (larger grains
are more rounded) phenoclasts of clinopyroxene (5 to 35%), plagioclase (albite,
15 to 34%) and rare pargasitic amphibole (0 to 8%); basaltic groundmass clasts
(with or without phenocrysts, 19 to 43%). Matrix formed by recrystallisation
and devitrification of finest fractions. Minor bedding and clast alignment;
erosional contacts between size fractions suggest numerous pulses of activity
rather than gravity settling; grain size gradational to basaltic breccia.

Very fine sand to cobble size (0.05mm to 10cm, rare boulders to 1m), very
poorly sorted, immature. Angular (larger clasts are more rounded) basaltic
lava cobbles and boulders; basaltic groundmass clasts (with or without
phenocrysts, 26 to 61%); phenoclasts of clinopyroxene (15 to 28%),

plagioclase (albite, 3 to 11%) and rare amphibole; limestone cobbles to 15cm
and calcite grains to 3mm. Matrix recrystallised and devitrified material.

Rare calcite cement. Larger limestone component (cobbles and grains) than
basaltic arenite; clast alignment indicates minor current activity; slump
deposit emphasised by sorting and clast size decrease away from larger clasts.

Porphyritic with phenocrysts of subhedral to euhedral plagioclase (albite,
glomeroporphyritic, 25 to 35%) to Smm long, and rare subhedral
$i05,=53-57%)| clinopyroxene (0 to 9%). Groundmass (50 to 58%) pilotaxitic, of albite laths
- with interstitial chlorite and sphene. Albite partially altered to sericite,
chlorite and rare calcite. Clinopyroxene grossly altered to calcite and sphene.
Original groundmass partly glassy; devitrified and now mainly chlorite and
sphene. Porphyroblastic aggregates of calcite, epidote, chlorite and prehnite.

andesitic Very fine to coarse sand size (0.01 to 3mm), moderately sorted-ash fraction
arenite to very poorly sorted in coarser rocks, immature. Angular (larger clasts
are more rounded) phenoclasts of plagioclase (albite, 34 to 43%) and rare
clinopyroxene; andesitic groundmass (with or without phenocrysts, 36 to 55%).
Matrix formed by recrystallisation and devitrification of finest fractions; rich

in chlorite and sphene. Minor bedding, clast alignment and size grading indicate
some current activity.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 175

Petrography

The Cudgegong Volcanics consist of lavas of basaltic, basaltic andesite and
andesitic composition together with associated very fine- to coarse-grained arenite and
breccia (Table 1). The rocks show the imprint of prehnite-pumpellyite to greenschist
facies metamorphism (Offler and Pemberton, 1983).

Fauna, Age and Correlation

The only recognized fossils in the Cudgegong Volcanics are found in a thin marl,
up to 20cm thick, underlying the basaltic breccia with limestone clasts marker horizon
in the central area (Figs 3 and 4; GR 568733). Pickett (1982a) identified the coral Plasmo-
porella sp. and the alga Vermzporella sp. from autochthonous limestone beds in the marl.
Pickett (1978) reported the same coral and alga, together with a Gisbornian conodont
fauna, from limestone clasts in a breccia towards the top of the Sofala Volcanics.

A 500 15 30, 300 40 20 30 350 300 B

Basaltic lava, fine- to Finely bedded Basaltic lava
coarse-grained arenite, basaltic ash and breccia
and breccia

Other symbols as for Fig. 3. * Fossil locality in marl
Distances in metres; not drawn to scale
Fig. 4. Representative cross-section through the Cudgegong Volcanics.

The Cudgegong Volcanics are here correlated with the upper parts of the Sofala
Volcanics on the basis of similarities in lithologies (Packham, 1968; Barron, 1976;
Gilfillan, 1976), clinopyroxene chemistry (Pemberton and Offler, 1985) and fauna. The
fauna occurs towards the south margin of the Cudgegong Volcanics where the strata are
southwest dipping and apparently upright, and as this suggests the fauna occurs towards
the top of the unit, a Gisbornian age is indicated for this part of the Cudgegong
Volcanics.

Structure

The Cudgegong Volcanics in the central, northwest and southeast areas (III, I and
IV respectively in Fig. 3) form the core of a northwest-trending anticline, with an over-
turned northeast limb. The southwest area (II) is considered part of a southwest-dipping
Ordovician-Silurian block faulted into position against Devonian rocks.

Folding within the Volcanics cannot be recognized. Marker horizons, where
present, are traceable only over short distances yet they strike parallel to the contact with
the Willow Glen Formation and do not indicate any sequence repetition. In the central
area (III), the sequence is apparently simply southwest dipping (data from finely lami-
nated ash, flat bedded arenite and crudely aligned clasts in breccia) and younging, from

PROC. LINN. SOC. N.S.W., 111 (3), 1989

176 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

the reworked top of an andesitic lava, indicates strata, near the south margin at least, are
upright. Possible internal folding hinders attempts to produce a representative section
for the Volcanics and as a compromise, a representative cross-section has been compiled
for the northwest of the central area (Fig. 4).

Environment of Deposition

The majority of rocks in the Cudgegong Volcanics are immature, grain-supported
arenite containing poorly sorted, randomly-oriented angular clasts of their respective
lavas forming thick apparently structureless outcrops. The arenites satisfy many of Cas
and Wright’s (1987) criteria for formation by volcaniclastic debris flow.

The majority of arenites were derived from an andesitic parent; however, the
outcrops display no features indicating possible source areas. Exposures of basaltic
rocks are limited, yet in the northwest of the central area (III), a possible near-vent
location is suggested by outcrops of basaltic breccia with limestone clasts, andesitic and
basaltic lava and possible syenite sills which fulfil some of the required near-vent criteria
(Williams and McBirney, 1979; Cas and Wright, 1987).

Preservation of limestone clasts in breccia together with the underlying marl in the
northwest of the central area (III), calc-silicate and chert horizons in the southeast area
(IV), and the identification of probable pillow lava (GR 619696 and GR 580747)
indicate a marine environment for both the volcanic and quiescent periods. The
autochthonous fauna in the marl indicate that in some areas, at least, shallow marine
conditions prevailed.

The data suggest the Cudgegong Volcanics formed by slumping of unstable
andesitic and basaltic debris from volcanic island slopes in a subaqueous environment.

WILLOW GLEN FORMATION (Pemberton 1980a)

The Willow Glen Formation, named after the former Willow Glen property 3km
southeast of Cudgegong, was initially described and named by Pemberton (1980a). The
extensive exposure of the unit was noted by Pemberton (1980b) and in this paper the
previous description of the unit is reviewed and expanded.

Stratigraphic Relationships

The formation crops out in a number of separate areas (Fig. 1; areas (a) to (f) in
Fig. 5). In each area, the unit is conformably overlain by the Windamere Volcanics
(contact clearly exposed in the new Cudgegong-Mudgee road at GR 575748); the
exception is the southeast area (f) where it is overlain conformably by the ‘loolamanang
Formation.

The formation consists of conglomerate (dominated by silicic volcanic clasts)
grading to pebbly litharenite and litharenite, shale, fossiliferous limestone, and rare
rhyolite horizons (Table 2).

In the southeast area (f), Pemberton (1980a) established a sequence of basal
litharenite, lower limestone bed (biomicrite with silty interbeds), massive dacitic ash
grading to well-bedded shale, and upper limestone bed (biomicrite with abundant silicic
volcanic clasts and common pentamerid brachiopods).

To the northwest, the formation differs markedly with the introduction of abundant
lensoidal conglomerate horizons and thinly bedded shale and arenite with common
limestone lenses. The most complete and representative exposure of these rocks occurs
in the eastern portion of the central area (e from Fig. 5). Unfortunately these exposures
lie close to the Cudgegong River and now the majority of these, together with several
important fossil localities, are covered by the waters of Windamere Dam. Rocks
typically exhibit rapid lateral facies change which produces considerable variation 1n

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 177

willow) :
Bend

WINDAMERE B Representative
AM 5

D,
section
18 cal [= | Windamere Volcanics

Willow Glen Formation

conglomerate ill limestone
Outcrop areas a to f for

rhyolite ae no outcrop
the Willow Glen Formation

shale, arenite, pebbly arenite

symbols as for Fig. 1

[=| Cudgegong Volcanics

Fig. 5. Geology of the eastern portion of the central area (e) of the Willow Glen Formation.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

178 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

both thickness and extent of the outcrops. Consequently the prominent conglomerate
horizons, within the recessive shale, arenite and limestone, cannot usually be traced as
marker horizons for distances greater than several hundred metres.

TABLE 2
Petrography of the main rock types in the Willow Glen Formation

conglomerate | Grey/white to red, very fine sand to cobble size (4cm), very poorly sorted,
immature. Subrounded to rounded white and black silicic volcanic clasts (80%),
fragmental plagioclase and quartz grains to 3mm, and rare shale, arenite and
limestone clasts. Matrix of fragmental quartz and plagioclase; strongly
recrystallised. Very fine siliceous cement, commonly with iron oxide or
chlorite. Gradational to pebbly arenite. Plagioclase grains altered to
sericite and calcite; matrix texture obscured by recrystallisation and
abundant secondary calcite veining.

litharenite Grey/brown to red, very fine-— to coarse-grained (2mm), moderately to poorly
sorted, immature. Angular to subrounded (larger grains are more rounded)
silicic volcanic, plagioclase, quartz and calcite grains with rare shale clasts.
Matrix strongly recrystallised; cement siliceous with common secondary iron
oxide and chlorite. Common secondary calcite veins and irregular-shaped
aggregates mask much of texture.

Grey/brown, very fine-grained siliceous material with rare angular quartz
and plagioclase grains to 0.1mm. Abundant aligned secondary white mica.

limestone Unsorted biomicrite and biosparite, and intrasparite; all with.common
detrital silicic volcanic clasts, and quartz and plagioclase grains. Silicic
volcanic pebbles (3cm) abundant in upper limestone bed of southeast outcrop
area. Jones et al. (1987) report oolitic limestone (intrasparite) from
GR 652679.

Stratigraphic sections from all outcrop areas are included in Fig. 6. The thickness
of section 2 may be exaggerated as the sequence through area (e) is probably repeated by
folding. Comparison of these sections has emphasized a number of similarities and
differences.

First, the majority of the Willow Glen Formation consists of lensoidal conglomer-
ate, arenite and pebbly arenite horizons within a fossiliferous sequence of thinly bedded
shale and arenite with limestone lenses. The two obviously different areas are: the
southeast area (f; section 1 of Fig. 6) where conglomerate is absent and shale with lime-
stone lenses are sparse; and at the northwest of area (d) (section 6 of Fig. 6) which is
dominated by thick lensoidal conglomerate.

Second, the distinctive pebbly limestone (abundant silicic volcanic clasts and com-
mon pentamerid brachiopods) of the upper limestone bed, southeast area (f) also occurs
in the limestone horizon in the southwest area (b); the southeast limestone bed of area
(c) (GR 612698); and numerous limestone lenses in the eastern portion of the central
area (e). Similar limestones are not recorded in the northwest of the district.

Third, the thickness of the sections and the outcrop width of the areas varies
dramatically and this suggests that the top of the formation was not a flat topographic
surface, rather that the essentially conformable Windamere Volcanics filled erosional or
nondepositional depressions on this surface.

Fauna, Age and Correlation
Numerous fossil localities have been recorded from shales (brachiopods and trilo-
bites) and limestones (brachiopods and corals) in the Willow Glen Formation (faunal

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 179

area d s oy. Location of stratigraphic sections
poaicutting i _ . for the Willow Glen Formation

symbols as for Fig. 1

km
ia
a N

WINDAMERE

? faulted

Willow Glen Formation

conglomerate

limestone
aX | Windamere Volcanics
litharenite
shale A Toolamanang Formation
pebbly litharenite
o!
a Cudgegong Volcanics faulted

shale with limestone lenses

shale with arenite interbeds

Oolitic limestone

dacitic ash

HERB I

Fig. 6. Representative stratigraphic sections for the outcrop areas of the Willow Glen Formation. No section
from the northwest area.

PROG. LINN. SOG. N.S.W., 111 (3), 1989

180 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

lists in Appendix). Pickett (1982a) proposed a Wenlockian to Ludlovian age based on the
identification of the brachiopod Kzrkidium and the corals Phaulactis, Halysites ortho-
pteroides, Desmidopora multitabulata, and Pycnostylus scalariformis from the limestone
localities. Recovery of conodonts from limestone samples was low with the best-
preserved specimens being referable to Ozarkodina ranuliformis, a species considered to
occur over a broad Silurian age in Australia (Pickett, 1982a).

Strusz (pers. comm., 1984) recognized that the Willow Glen Formation shale fauna
was similar to part of the fauna from Coppin’s Crossing, near Canberra (Strusz, 1982).
He identified the brachiopods Salopina ?mediocostata, Aegiria cf. norvegica, Morinorhynchus
oepiki and Coelospira cavata, together with Maoristrophia (Strusz, 1983) and the trilobite En-
crinurus mitchellr (Strusz, 1980). He suggested a Late Wenlockian to Early Ludlovian age
for the fauna.

Silurian sedimentary sequences unconformably overlie Sofala Volcanics equiva-
lents at at least four known localities: Sofala and east of Wattle Flat (Packham, 1968);
Palmers Oakey (Powell, 1984, based on Bischoff and Fergusson, 1982) and now at
Cudgegong. In the Sofala-Wattle Flat area, Packham (1968) identified a coral-trilobite-
brachiopod fauna, with sparse graptolites, from the Tanwarra Shale. Based on tentative
identification of the graptolites, he suggested the Tanwarra Shale may be as old as Late
Llandovery. However, in the Tanwarra Shale equivalents at Palmers Oakey, Bischoff and
Fergusson (1982) recognized a very Late Wenlockian age for an extensive conodont
fauna. The Late Wenlockian to Early Ludlovian age proposed for the Willow Glen
Formation, after Pickett’s and Strusz’s faunal identifications, 1s consistent with the age of
the strata at Palmers Oakey and perhaps the age of the Tanwarra Shale at Sofala.

Structure

The southeast, northwest and central areas (f; a; and c, d and e respectively from
Fig. 5) lie on the limbs of an anticline whereas the southwest area (b) is part of an
Ordovician-Silurian block faulted against Early Devonian strata.

In both the southeast and northwest, bedding data are limited and mapping of
formation boundaries and age relationships indicate the anticlinal structure. Bedding
data from the upright southwest-dipping area (c) have a mean limb dip of 35° with a
modal strike of 315° whereas data from the southwest-dipping overturned (facing from
cross-bedded pebbly arenite) areas (d) and (e) have a mean dip of 75° with a modal
strike varying from 310° to 320°. Combined limb data indicate a near-horizontal fold
plunge with both southeast and northwest plunge components. Within-limb parasitic
folding is common, with similar fold styles and plunge to the large scale structure.

Environment of Deposition
Conglomerate, arenite and pebbly arenite horizons

These horizons have many features indicative of a fluvial channel-fill origin. These
include the very poorly-sorted yet well-rounded nature of the clasts; common interbed-
ding of lithologies producing flat lamination, with clast alignment in the coarser beds;
cross-bedding with pebbles aligned parallel to, and with fining upwards sequences in,
the cross beds as well as low angle beds asymptotic to the cross-bed base; and their
lensoidal shape. The lenses vary greatly in thickness (a few metres up to 200m) and
lateral extent (10m up to several km) providing evidence for an extensive channel system
preserving single channel-fill events (thickness in the order of 5m) as well as multiple
channel-fill build-up.

Shale and arenite with limestone lenses

Strusz (pers. comm., 1984) noted the faunal and lithological similarity of these

PROC. LINN. SOC. N.S.W,, 111 (3), 1989

J. W. PEMBERTON 181

rocks to the strata near Coppin’s Crossing, Canberra (Strusz, 1982). He considered the
latter to represent Boucot’s (1975) benthic assemblage 3 — the subtidal zone with both
quiet and rough water reef communities below 6m water depth. The thinly bedded
fossiliferous shale, arenite and zn sztu limestone of the Willow Glen Formation represent
a dominantly quiet water, low energy environment. The thicker lensoidal limestones
may indicate local mounds with rough water conditions, as indicated by the
pentamerid-coral fauna (Boucot, 1975), and provide the higher velocity currents
necessary to transport and deposit the silicic volcanic pebbles typical of some of these
limestones.

The detailed study of the strata at Willow Bend (GR 652681 from Fig. 5; Jones e¢
al., 1987) demonstrates a thin clastic regressive unit within open-marine limestone. The
basal shale (low energy subtidal environment below storm wave base) is sharply overlain
by a very shallow marine oolitic limestone which was later subjected to evaporation on
supratidal flats. The sequence was then partly eroded by fluvial scours prior to trans-
gression with initially another localized oolitic limestone and then by more open-marine
limestone.

In summary, the majority of the formation represents deposition on subtidal to
supratidal flats affected by transgressive/regressive cycles. The overall pattern is one of
fluctuating sea level and energy regimes with localized rough water limestone mounds
surrounded by quiet water deposition. The inferred flats were incised by fluvial
channels accompanying periods of sea-level regression.

Variations

The northwest of area (d) and the southeast area (f) are lithologically different from
the majority of the formation. In the former area, thick conglomeratic horizons
represent substantial fluvial channel-fill accumulations possibly providing the source
direction for the channel system incising the tidal flats to the southeast and southwest.
By comparison, in the southeast area, the absence of conglomeratic horizons, and the
thick ash/shale and arenite beds with poorly fossiliferous limestone suggest open-marine
conditions to the southeast of the tidal flats.

Depositional setting

Comparison of the Willow Glen Formation from the northwest to the southeast of
the district indicates a typical 20-25km wide coastal zone, similar to present-day
examples along the north and northeast Australian coast, with progression from the
fluvial channel zone through a tidal flat to more open-marine conditions.

WINDAMERE VOLCANICS (Pemberton 1980b)

These rocks were included by Offenberg et al. (1971) in the Sofala Volcanics.
However, recognition of their Late Silurian age and dacitic nature led Pemberton
(1980b) to propose the Windamere Volcanics, named after the former property
upstream from the Windamere Dam wall.

Stratigraphic Relationships

The formation crops out to the northwest of Cudgegong initially as a 1km-wide belt
which widens northwest of Windamere Dam reaching a maximum width of 4km in the
Millsville area (Fig. 1). These exposures lie on the limbs of the major northwest-trending
anticline and have been described (data summarized in Table 3) as the northern limb

(Fig. 7) and southern limb (Fig. 8).

PROC. LINN. SOG. N.S.W., 111 (3), 1989

182 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

symbols as

for Fig. 1

Location of the
“northern limb” (Fig. 7)
es Windamere
Dam wall and “southern limb’

(Fig. 8) of the

Windamere Volcanics

Late Devonian

Early Devonian

Willow Glen Formation /

Cudgegong Volcanics

Windamere Volcanics

dacite lava and breccia

rhyolite

dacitic conglomerate

no outcrop

localities referred to in Table 3

homestead

F fault

Fig. 7. Lithological variation for the northern limb of the Windamere Volcanics.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

183

J. W. PEMBERTON

Permian conglomerate

Millsville Formation

Windamere Volcanics

Horse Flat conglomerate

c
°
N

=
°

£
£
=
)
Gy

U
)

x
ra
fe)
=
Go)
L

Ey Le]

other symbols as for Fig. 7

Fig. 8. Lithological variation for the southern limb of the Windamere Volcanics.

PROC. LINN. SOG. N.S.W., 111 (3), 1989

184

Petrography

CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

The Windamere Volcanics are characterized by thick and extensive exposures of
undifferentiated dacite lava and breccia, with common rhyolite horizons at all strati-
graphic levels (Table 4). Less common rock types include dacitic arenite (with sparse
pebbles) to rare ash-size rocks; dacitic conglomerate; dacitic breccia with limestone
clasts; flow layered dacite lava; and shale, with rare limestone clasts. Positions in the
sequence and best exposures are included in Table 3.

Northern
limb

Middle
sequence

Upper
sequence

Rhyolite

TABLE 3

Extent of and sequence within, the Windamere Volcanics Extent

Overturned limb bounded by unconformity with Late Devonian rocks along
Cudgegong River valley and to southeast by Cudgegong Fault. Unit top not
exposed yet conformably underlain by Willow Glen Formation over outcrop length.

Provides more complete sequence with both conformable underlying (Willow Glen

Formation) and overlying (Millsville Formation) contacts, the latter only in
certain northwest localities. The Millsville Formation and rhyolite of Windamere
Volcanics are both disconformable and faulted with Early Devonian strata of
Queens Pinch Belt.

Sequence and best exposures

Dominated by dacite lava and breccia. Variations include: numerous conformable
rhyolite horizons, to several m thick, both at base and slightly higher in
sequence; in Ironstone Creek (GR 657692), Sm of dacitic arenite with limestone
and rhyolite clasts, and black shale, with limestone lenses, occur above an

initial 20m of dacite lava (contact not exposed); and on Horse Flat (GR 503810)
Willow Glen equivalents are sharply overlain by 100m of coarse conglomerate
(dacite and rhyolite clasts), pebbly arenite and arenite.

Dacite lava and breccia dominant (extensively exposed in new road cuttings and
dam spillway), with greater diversity of less common rock types. Include: dacite
lava with polygonal cooling joints; dacite lava with rounded mafic xenoliths to
several cm across; flow layered dacite lava; discontinuous dacitic conglomerate
beds; rare dacitic breccia with limestone clasts; coarse dacitic breccia with

flow layered clasts to 1m across (GR 522774); common thin conformable rhyolite
horizons; and 25m thick reworked dacite horizon (GR 528781) of thinnly bedded
arenite, shale with sandy interbeds (ripple marks), coarse breccia, conglomerate
with arenite beds (cross—beds and scour and fill structures). Facing from
sedimentary structures indicate horizon upright.

Proportion of detrital dacitic rocks increases within dominant dacite lava and
breccia. Detrital lithologies include: breccia with limestone clasts; arenite

and pebbly arenite; 100m thick shale bed (GR 518774).with rare thin limestone
and conglomerate lenses; and 10m thick dacite boulder horizon (GR 518771) only
a few m below conformably overlying Millsville Formation.

Flow layered lava, with rare breccia and conglomerate, occurs at all

stratigraphic levels. Thickness varies from few m to 100m, with thicker horizons
as prominent ridges. On the southern limb, one such 100m thick apparently
conformable body of flow layered lava forms prominent steeply southwest—dipping
scarp for 5.5km length. Best conformable contacts with dacite at GR S07777

and GR 630716.

Primary dacite lava is distinguished from the fragmental rocks by the lack of
fragmental phenocrysts (rounded, embayed quartz, subhedral to euhedral plagioclase
and amphibole); the glomeroporphyritic nature of phenocryst aggregates; the presence
of rare amygdales; and the clear distinction between phenocrysts and groundmass (15 to
35% phenocrysts with the maximum density 49%). However, recognition of primary

PROC. LINN. SOC. N.S.W,, 111 (3), 1989

J. W. PEMBERTON 185

textures is hindered by the effects of prehnite-pumpellyite to greenschist facies
metamorphism. Moreover, recrystallization of the groundmass produces granoblastic
quartz masses. Further, breakdown of the groundmass varies from the minor develop-
ment of fine-grained interstitial chlorite to the major development of distinct chlorite-
rich and quartz-rich portions, the latter producing a brecciated appearance. Another
feature hindering the recognition of primary textures, especially in rocks towards the
top of the formation, is the secondary silicification of highly fractured dacitic rocks and
the development of siliceous spherulitic concretions in numerous rhyolite horizons.

‘TABLE 4
Petrography of the main rock types in the Windamere Volcanics

dacite lava Porphyritic with phenocrysts of subhedral plagioclase (albite,
Si05,=65-69%| glomeroporphyritic, 11 to 32%) to Smm long; embayed quartz (fractured, 0 to
is 6%, fine-grained recrystallised margins) to 3mm across; and rare subhedral

to euhedral amphibole (hornblende, 0 to 9%, X=pale brown, Y=light brown,
Z=brown/green) to 1mm long. Fine-grained and recrystallised groundmass (51
to 87%) dominated by anhedral quartz and plagioclase. Rare ovoid amygdales
contain chlorite. Albitised plagioclase phenocrysts and groundmass grains
partially altered to chlorite, sericite, epidote, pumpellyite, prehnite and
calcite. Chlorite very common in groundmass (to 16%) as interstitial material
in less altered rocks, and as irregular-shaped masses, to several mm across,
and as pseudomorphs after plagioclase and amphibole in more altered rocks.
In the most altered rocks, groundmass segregates into quartz-rich and
chlorite-rich portions.

Very fine-grained to cobble size (0.0l1mm to 10cm; rare boulders to 1m),
very poorly sorted, immature. Angular (rounded larger clasts) dacite lava
cobbles and boulders, dacite groundmass clasts, plagioclase and quartz
phenoclasts. Matrix of finest sized dacite fragments (dominated by
recrystallised quartz and plagioclase). Rare primary calcite cement.
Alteration of phenocrysts and groundmass similar to that described for
dacite lava. Development of metamorphic textures, in particular breakdown
of groundmass, may hinder recognition of primary fragmental nature..

dacite Very fine-grained to very coarse-grained sand (0.01 to 3mm), poorly sorted,
arenite immature. Angular dacite groundmass clasts, quartz and plagioclase
phenoclasts, and rare dacite lava clasts. Matrix formed by recrystallisation
and devitrification of finest dacitic fraction. Alteration as described for
dacite lava and breccia. Sharp erosional contacts between varying grain sizes.

rhyolite lava | Sparsely porphyritic with phenocrysts of embayed quartz (finely
recrystallised margins) to 3mm across, and rare subhedral feldspar (strongly
altered, untwinned) to 5mm long. Coarsely recrystallised groundmass of
polygonal quartz and irregularly— shaped altered feldspar. Common flow
layering as compositional banding of quartz-rich and —poor horizons to Smm
_thick. Feldspar phenocrysts very strongly to completely altered to sericite
and chlorite; groundmass grains less severely sericitised. Spherical
siliceous concretions, to 4cm across, are very abundant towards the top of
the unit.

Age and Correlation

No fauna has, as yet, been found in the Windamere Volcanics. However, the unit is
considered Late Silurian as it is conformably underlain and overlain by Wenlockian to
Ludlovian strata — the Willow Glen Formation and the Millsville Formation. Rhyolite
overlying the latter is disconformably overlain by Early Devonian strata.

In the Sofala district, rhyolite lava, arenite and breccia (the Wenlockian Bells Creek
Volcanics of Packham, 1968) conformably overlies the Tanwarra Shale and 1s overlain

conformably by the deep water Chesleigh Formation deposited on the eastern margin of
the Hill End Trough.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

186 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

Structure

The outcrop areas of the Windamere Volcanics lie on the limbs of the major anti-
cline. On the southern limb, facing (from the reworked dacitic horizon and the confor-
mably overlying Millsville Formation) indicates the southwest-dipping sequence is
upright. A composite representative section (Fig. 9) has been compiled from the basal
contact with the Willow Glen Formation on the Peckham property (Fig. 8), incorporat-
ing a number of northeast-southwest traverses, to the uppermost rocks, including the
Millsville Formation, below the Early Devonian disconformity along Back Creek (GR
521766). The southwest-dipping strata of the northern limb are considered overturned,
based on the relationship with older units as well as cleavage vs bedding orientation.

; Mullamuddy Formation
Early Devonian

dis conformity

a
re)
=)
3

Millsville Formation

rhyolite

dacite lava

dacite arenite
with breccia
xenoliths

ET
TOT

black shale

flat and cross beds
ripple marks

dacite breccia
with breccia

horizons include
arenite, breccia,
conglomerate, shale

INS BROUE

flow layered
dacite lava

om 4 with limestone clasts

Willow Glen
Formation

Fig. 9. Composite representative section through the Windamere Volcanics.

Common northeast-trending faults clearly offset rhyolite horizons on the southern
limb (Fig. 8), as well as the Millsville Formation; the Cudgegong Volcanics-Willow Glen

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 187

Formation sequence; contacts with the Devonian strata and Devonian sequences in the
Queens Pinch Belt. A similar pattern of faults affects strata on the northern limb (Fig.
7). The displacement along the faults varies from small scale (several metres) up to
hundreds of metres. The faulting is approximately normal to the regional fold axis and
probably represents tear or compartmental faults (Davis, 1984) which formed during
the regional folding and acted as transverse strike-slip faults accommodating the defor-
mation of a thick rock mass.

Environment of Deposition

The eruptive environment, at least during periods of volcanic quiescence, is
considered to have been shallow marine to probably emergent. Evidence includes: shale,
with apparently zm sztu limestone lenses, at the base and top of the unit; shale, arenite and
breccia, all with limestone clasts; and traction current structures (in the reworked
horizon [GR 528781] ) indicating fluctuating energy conditions in a shallow marine
environment. In addition, the lensoidal conglomeratic body on Horse Flat (GR 510815,
Fig. 8) together with other common conglomeratic lenses (Table 3) suggest a fluvial
channel origin. Further, the underlying Willow Glen Formation was deposited in a
shallow marine to supratidal environment with regression to subaerial conditions and
the overlying Millsville Formation includes shallow marine and beach deposits.

The eruption of silicic lava into a shallow marine to subaerial environment should
produce dominantly pyroclastic detritus with short thick lava bodies of limited aerial
extent (Cas and Wright, 1987). However, the Windamere Volcanics are dominated by
dacite lava, with less common breccia and rhyolite horizons.

An explanation for the high volume of dacite lava could be emplacement as a lava
dome. The growth of silicic domed masses around vents, and shallow intrusions (crypto-
domes), is preceded by highly explosive activity followed by long periods of non-
explosive dome growth capable of producing thick lava masses. Recent subaerial
examples include activity at Mt St Helens (Swanson e¢ al., 1987), the islands of Lipari
and Vulcano (Sheridan et al., 1987) and the South Sister volcano (Scott, 1987). The
initial explosions decrease the volatiles available for later magmatic pulses thus provid-
ing the degassing mechanism for emplacement of thick non-explosive lava bodies (Fink
and Manley, 1987). However, Newhall and Melson (1983) record numerous examples of
post-dome explosions which activated partial dome collapse and led to autobrecciation
of parts of the lava body.

If the Windamere Volcanics were emplaced as a thick domal body, the initial
degassing mechanism may have produced the fragmental material of the Toolamanang
Volcanics, a lateral mass-flow equivalent of the Windamere Volcanics. The common
breccia may have formed from later explosive events causing autobrecciation by small
scale dome collapse. In this model the abundant conformable rhyolite horizons could be
either extrusive (with the thicker bodies towards the top of the volcanics representing
terminal activity) or emplaced as contemporaneous sills.

TTOOLAMANANG FORMATION (Pemberton 1980a)

The unit was named after the historic Toolamanang property, south and southwest
of Cudgegong. To the southeast of Cudgegong, Pemberton (1980a) recorded rhyodacite
lava, breccia and arenite (the Ioolamanang Volcanics); these rocks had been assigned by
Offenberg et al. (1971) to the Sofala Volcanics. Subsequent mapping to the southwest of
Cudgegong (Pemberton, 1980b) revealed much greater exposure of the unit with a lack
of primary volcanic features; thus the original description of the Toolamanang Volcanics
has been considerably modified.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

188 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

Stratigraphic Relationships

To the southeast of Cudgegong, the Ioolamanang Formation overlies the Willow
Glen Formation with apparent conformity (Pemberton, 1980a). However, to the north-
west along Limestone Creek valley, the unit is faulted against Ordovician and Silurian
rocks where the contact 1s marked by a prominent ferruginous zone tens of metres thick.

The unit crops out over 14km in length with a width up to 5km (Fig. 1). Near Mt
Bocoble, an apparently conformable overlying contact is exposed with limestone (with
pentamerid brachiopods and halysitid corals), breccia (limestone clasts in a sandy
matrix), conglomerate and shale which resemble, from the fauna and lithology, parts of
both the Willow Glen Formation and Millsville Formation. To the southeast of Mt
Bocoble, this contact is poorly exposed and the ?overlying rocks are discontinuous
unfossiliferous limestone beds.

Lithologies

The Toolamanang Formation is a massive, structureless succession of fine- to
coarse-grained arenite with common black mud horizons and sporadic basaltic out-
crops. The main rock types consist of dacitic detritus as lithic arkose and feldspathic
litharenite; with common very fine ash size rocks and fine-grained breccia. The rocks
are texturally similar, being very poorly sorted with angular plagioclase (albite), less
common quartz and rare hornblende fragments, up to 3mm across, and a highly vari-
able content of dacite groundmass clasts ranging from several mm to 3cm across. Clasts
are tightly packed, with a recrystallized matrix of fine dacitic detritus. Lithological
variation in the succession, in which no sequence or marker horizons could be estab-
lished, is minor and involves rapidly varying grain sizes. Basaltic blocks, petrographi-
cally and chemically similar to basaltic rocks in the Cudgegong Volcanics (Pemberton
and Offler, 1985) occur throughout the unit, varying in size from tens of centimetres up
to 3m across.

The best exposures occur in several cuttings on the new Cudgegong-Mudgee road
(GR 631661 to GR 618681) and their associated drill cores. They are dominated by fine-
to coarse-grained arenite with massive black mud (fine-grained dacitic ash) beds, from
several cm to Im thick. The bases of the sandy horizons show loading and slumping of
the sand squeezed and injected into the mud whereas the tops are generally flat and
sharp. Several of the basal zones, which vary from a few cm to Im thick, have a dis-
oriented fabric of irregularly-shaped sandy masses representing intensely churned and
squeezed sand within the mud layers. Fine-grained breccia patches (with randomly-
aligned arenite and black mud clasts) and angular and rounded basaltic blocks are
common.

Both the cuttings and the cores show: firstly, that the basaltic material is fragmen-
tal, varying greatly in size and that this material has been deposited in the muddy and
sandy detritus; and secondly, that the grain size of, and thickness of the units within, the
dacitic detritus varies rapidly, indicating multiple depositional episodes, and that
contacts between units are sharp and show evidence of loading and slumping.

Age and Correlation

No fauna has, as yet, been found in the Toolamanang Formation. The unit 1s
considered Late Silurian on the basis of the Wenlockian to Ludlovian age of the confor-
mably underlying Willow Glen Formation and the preliminary dating of the fauna from
?overlying limestone near Mt Bocoble as Late Silurian.

Lithologies in the formation consist principally of volcanic detritus of similar
composition to dacite lava, the dominant rock type in the Windamere Volcanics.
Together with similarities in age and underlying and overlying formations, the

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 189

Toolamangang Formation 1s considered a lateral equivalent, and fragmental version, of
the Windamere Volcanics.

Correlatives of the Windamere Volcanics, the Bells Creek Volcanics (Packham,
1968) and the Mullions Range Volcanics on the Molong High (Hilyard, 1981); both
contain a high proportion of fragmental rhyolitic and dacitic material as does the
Toolamanang Formation.

Structure

Rare bedding data display the regional southeast strike with steep southwest dips.
R. Offler (pers. comm.), in a study to the northeast of the new Cudgegong-Mudgee road
(Fig. 1), confirmed the southwest-dipping trend yet he noted dip reversals within and
between outcrops indicating small scale southeast-plunging folds.

There has been no attempt to compile a representative section due to the lack of
established sequence, the unknown internal structure and the poorly known southern
portion of the formation. If the apparently overlying strata are Millsville Formation
equivalents, the formation may be simply dipping to the southwest and the 5km outcrop
width produces a thickness of approximately 3.5km. Alternatively, the outcrop pattern
near Cudgegong suggests a synclinal structure (Pemberton, 1980a) with an upright
sequence on the eastern limb and, if the ?overlying rocks are Willow Glen Formation
equivalents, the larger scale structure may be a northwest-plunging syncline with the
southwest limb overturned. As a consequence the unit thickness may be nearer 1.5 to
2km.

Environment of Deposition

The pervasive arenite of the Toolamanang Formation consists of angular, poorly
sorted, ash- to fine lapilli-sized dacitic detritus in massive structureless outcrops and
suggests deposition as an unsorted crystal and lithic ash formed from dense, gravity-
driven, volcaniclastic flows (Cas and Wright, 1987).

The black muddy rocks are composed of very fine-grained dacitic detritus and vary
from massive bodies to intensely-churned zones formed by slumping and loading of the
overlying sands. The rocks suggest deposition as mud flows associated with the coarser
ash-flow events. The black mud clasts in the breccia indicate transport of partially
lithified mud.

The basaltic rocks are considered part of the Cudgegong Volcanics basement which
has been included in the debris flows by erosion, or possibly explosive ejection, from the
flanks of the Late Silurian volcano and subsequently transported, with minor rework-
ing, prior to the dumping of the load into the sandy and muddy strata.

There are no direct indicators of a subaqueous or subaerial environment for the
debris flows, although the underlying, tentatively overlying and laterally equivalent
units exhibit shallow marine to emergent characteristics. The slumping so typical of the
arenaceous rocks can occur on fairly gentle slopes and this suggests the detritus was
deposited into slightly deeper water conditions than that of the northwest source.

It has been previously implied that the processes which initiated the debris flows of
the Toolamanang Formation may have provided the mechanism to erupt the thick
Windamere Volcanics lava body. The emplacement of similar silicic bodies produces a
variable yet significant volume of pyroclastic material, which acts as the explosive
degassing mechanism preceding thick lava growth (Heiken and Wohletz, 1987; Scott,
1987). There is no evidence of explosive activity preserved in the Toolamanang For-
mation; however, the variable grain size within the thick flow succession clearly
represents a large number of flow episodes. It remains to be proven that the episodes

PROC. LINN. SOC. N.S.W., 111 (3), 1989

190 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

were explosively initiated thus providing the volatile release necessary to allow the
Windamere Volcanics emplacement.

MILLSVILLE FORMATION (Powis 1975)

Offenberg et al. (1971) included these rocks in the Siluro-Devonian Gulgamree
Beds. However, Powis (1975) recognized their Late Silurian age and demonstrated map-
pable differences from the nearby Early Devonian sequences. He proposed the name
Millsville Beds, after the nearby property. Further mapping by Pemberton (1980b, and
more recently) has confirmed Powis’ ideas and herein the sequence is formally named
the Millsville Formation.

Stratigraphic Relationships

There are two main outcrop areas of the Millsville Formation here described as the
southeast and northwest exposures (Fig. 10; location from Figs 1 and 8).

The major rock type is breccia with dominantly limestone and minor dacite and
rhyolite clasts in a dacitic matrix. Subordinate rocks include: limestone (biosparite and
biomicrite); a gradational sequence of shale, arenite and fine-grained conglomerate of
calcareous and dacitic detritus; and well-defined dacitic conglomerate and breccia
horizons (Table 5). In addition, thick rhyolite lava, with rare breccia, occurs at the top of
the formation.

In both areas, the formation conformably overlies dacite of the Windamere
Volcanics, with gradational contacts recognized by an increase in fragmental dacitic
material and the appearance of limestone lenses and detritus. However, the basal rocks
in certain localities in the northwest area are discontinuous (up to 100m long) dacitic
conglomerate lenses up to 10m thick (Fig. 10).

The pervasive breccia is present throughout the formation with little lithological
variation. Limestone horizons vary from rare thin lenses, from several cm to 1m thick, to
several prominent zn situ bodies up to 300m long with thickness to 100m (Fig. 10).
Thinly-bedded fossiliferous shale, and flat-bedded arenite to fine-grained conglomerate
occur as discrete horizons or as gradational lenses within the breccia succession.

The northwest area and certain localities in the southeast area are conformably
overlain by a persistent rhyolitic succession. The rhyolite and remaining Millsville For-
mation are disconformably overlain by the Early Devonian Mullamuddy Formation
(Fig. 10).

In general, the formation is typified by rapid facies change. Marker horizons are
few and include the basal dacitic conglomerate in the northwest area and a prominent 10
to 15m thick dacite breccia horizon, outcropping for over 1.5km length, towards the
middle of the southeast area sequence (Fig. 10).

Fauna, Age and Correlation

Powis (1975) reported four fossil localities (Fig. 10), in limestone and shale, to which
he assigned a Silurian age. His faunal lists included: limestone — Phaulactis, heliolitids,
favositids, stromatoporoids, and pentamerid brachiopods; shale — Halysites cf. bellensis,
Encrinurus, Rhizophyllum, Eospirifer, Leptaena together with orthid, atrypid and rhyn-
chonellid brachiopods. Pickett (1982a) noted Kzirkidium, Thamnopora, Amphipora, and
?Propora, together with rare conodonts (not age-diagnostic), within the prominent lime-
stone of the northwest area. The fauna is clearly Silurian and the recognition of faunal
similarities with the Willow Glen Formation, together with the relationship of the for-
mation to the underlying Windamere Volcanics and the overlying Lochkovian to

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 191

Pragian Mullamuddy Formation, indicates a Late Silurian (Wenlockian to Ludlovian)
age for the Millsville Formation.

southeast exposures

breccia with
limestone and dacitic arenite [So] Mullamuddy Formation

dacite clasts

limestone Fed Early Devonian shales

Windamere Volcanics

dacite breccia dacite A rhyolite
conglomerate

fossil locality

Fig. 10. Geology of the two outcrop areas of the Millsville Formation.

The Ordovician-Silurian sequences at Cudgegong and Sofala are similar; however,
Windamere Volcanics-Ioolamanang Formation equivalents in the Sofala area are
conformably overlain by deep water rocks of the Hill End Trough sequence (Packham,
1968). However, in the Cudgegong-Mudgee district, Late Silurian rocks overlying silicic
volcanic sequences (the Millsville Formation) are deposited in a shallow marine
environment.

PROG. LINN. SOC. N.S.W., 111 (3), 1989

192 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

TABLE 5
Petrography of the rock types in the Millsville Formation

breccia Grey to pink, very fine sand to boulder size (1m), very poorly sorted,
immature. Angular to subangular (larger clasts more rounded) dominantly
limestone (poorly washed biosparite with limy mud fraction) clasts, from
Sem to 1m across, with subordinate dacite (lava and arenite), rhyolite
lava, shale, and siliceous clasts to 10cm across. Matrix supported; matrix
consists of fine-— to coarse-grained arenaceous dacitic and calcareous
detritus.

limestone Grey, cream to red; poorly washed biosparite (rich in corals, brachiopods
and stromatoporoids) with limy mud horizons; sparse to packed biomicrite;
and calcirudite patches (Powis, 1975).

dacitic Grey, fine sand to boulder size (0.5m), very poorly sorted. Rounded to

conglomerate angular dacite lava clasts, from 10cm to 30cm across, with rare boulders
to 0.5m. Clast-supported with minor dacitic matrix. No clast alignment.
Boulder horizon in southeast exposures similar texture yet clasts angular
and coarser.

dacitic Grey to brown, fine sand to boulder size (0.5m), very poorly sorted,

breccia immature. Angular dacite (lava and arenite) clasts from several cm to
20cm across, rarely to 0.5m; with common limestone clasts to several cm
across. Matrix supported; matrix of dacitic detritus.

Structure

There is no evidence of folding within the formation. Limited bedding data
indicate a consistently southwest-dipping unit up to 220m thick. Further, geopetals
(brachiopod and stromatoporoid orientations) in the limestone indicate upright
horizons. Consequently four representative sections (Fig. 11, locations from Fig. 10)
have been produced for areas of best exposure. Horizons within each outcrop area may
be crudely correlated; however, it is not possible to correlate between outcrop areas.

Environment of Deposition

The appearance of limestone and shale horizons at the top of the Windamere
Volcanics signals the start of volcanic quiescence in a shallow marine environment. The
basal dacitic conglomerate does not show features indicative of a fluvial channel origin
and the very poorly sorted, clast-supported lenses are likely to represent beach deposits.

The autochthonous limestone bodies contain a distinctive pentamerid brachiopod-
coral fauna typical of rough water limestone bank or mound communities (Boucot,
1975) on a shallow marine shelf. The shale horizons (in situ coral growth and a low
degree of brachiopod and trilobite fragmentation) formed during quiet water periods
and together with the limestone represent bank to lagoonal deposition.

The pervasive breccia suggests collapse of the limestone banks, possibly due to
storm activity, and the subsequent dumping of this material into dacitic detritus in
slightly deeper water.

The Millsville Formation represents deposition on a shallow marine shelf where
initially extensive bank to lagoonal areas, with minor beach deposits, were subsequently
affected by limestone bank collapse into slightly deeper water.

Persistent volcanism was then resumed as thick rhyolite masses conformably overlie
the shelf deposits, especially in the northwest area.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 193

southeast exposures northwest exposures

fine-grained conglomerate
with chert and limestone clasts

other symbols as for Fig. 10

Fig. 11. Representative sections through the Millsville Formation.

DISCUSSION

Nature of the Contacts with the Overlying Strata

Wright (1966) recognized that the Ordovician-Silurian sequences, described in this
paper, separated two belts of Devonian rocks — the Queens Pinch Belt and the Mt
Frome to Kandos Belt (Fig. 12).

Queens Pinch Belt (Carne and Jones 1919)

The belt consists of a number of fault-bounded slices of Early Devonian shelf and
turbidite facies rocks deposited on the western margin of the Capertee High (Wright,
pers. comm. and in Strusz, 1972). Contacts with the Ordovician-Silurian rocks are
mainly faulted, particularly in the Limestone Creek area; however, disconformable
contacts have been identified between the oldest unit in the Queens Pinch Belt, the
Lochkovian to Pragian Mullamuddy Formation, and the Wenlockian to Ludlovian
units — the Windamere Volcanics near Mt Margaret (GR 545757) and the Millsville
Formation along Back Creek (GR 520765).

Immediately north of Mt Margaret, the top of the Late Silurian rhyolite has been
reworked and at the base of the Mullamuddy Formation, a black shale, up to 1.5m thick,

PROG. LINN. SOG. N.S.W,, 111 (3), 1989

194 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

containing abundant rhyolite clasts grades up to an in situ limestone horizon. Near Back
Creek, the base of the Mullamuddy Formation has eroded deeply into the Millsville
Formation as the contact cuts through rhyolite and breccia successions as well as the
prominent dacite breccia horizon (Fig. 10). Here, the Mullamuddy Formation is a
breccia of unsorted rhyolite and limestone cobbles and boulders.

PERMIAN Shoalhaven Group

@ MT BOCOBLE
CARBONIFEROUS Aarons Pass Granite

LATE

MIDDLE

DEVONIAN

SILURO- Hill End Trough
DEVONIAN Sequences
eal ORDOVICIAN- SILURIAN

Fig. 12. Post-Silurian geology of the Cudgegong-Mudgee district.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON 195

Mt Frome to Kandos Belt
The belt comprises a thick sequence of Late Devonian fluviatile and shallow
marine successions (Wright, 1966) together with Early Devonian shallow marine

sediments and volcanics in the Cudgegong-Rylstone district (Pemberton, 1977;
Campbell, 1980; Millsteed, 1985; Cook, 1988; Colquhoun, 1989).

Cudgegong Fault (Game 1935)

Game (1935) proposed a large scale strike-slip fault (the Cudgegong Fault)
separated Ordovician-Silurian and Devonian sequences along the Cudgegong River
valley and to the southeast of Cudgegong. Pemberton (1980a) and Millsteed (1985) have
confirmed a faulted contact between Late Silurian and Early Devonian strata from the
Permian plateau to the headwaters of Ironstone Creek (Figs 5 and 12). Evidence for the
Cudgegong Fault in this area includes the truncation of folded Early Devonian units and
several members of the Late Silurian sequence. The fault plane is marked by a thick
linear ferruginous zone and a restricted fault breccia. In addition, C. L. Fergusson
(pers. comm., 1987) noted the development of a prominent zone of tectonic melange ina
road cutting (GR 652653) on the Cudgegong-Rylstone road along the line of the fault.

Lambian Unconformity (Powell and Edgecombe 1978)

A low angle discordance between the Late Devonian Lambie Group and a variety
of older rocks, the Lambian Unconformity, has been recorded from numerous localities
in the northeast Lachlan Fold Belt. In the Mudgee district (Fig. 12), the basal Late
Devonian unit, the Buckaroo Conglomerate of Wright (1966), crops out continuously
from Mt Frome to the Carwell Creek district.

Near Mt Frome, Powell and Edgecombe (1978) record an angular discordance of
5° to 24° where the Buckaroo Conglomerate overlies the latest Early to early Middle
Devonian sequence (Garratt and Wright, 1988). To the southeast, the contact is covered
by the Cudgegong River floodplain. An exception exists near the Windamere Dam
spillway where the Lambian Unconformity can be recognized along the road from the
spillway to the observation deck. Here the top of the Windamere Volcanics is
represented by 5m of intensely weathered dacite in sharp contact with the northeast-
dipping Buckaroo Conglomerate. Gross (1982) noted a discordance of up to 28° with the
Buckaroo Conglomerate.

To the south of Cudgegong, the Ordovician-Silurian strata are intruded by the
Middle Carboniferous (320 Ma, Vickary, 1983) Aarons Pass Granite, a massive biotite
granite/adamellite stock of 10km diameter. The strata are unconformably overlain by
thin flat-lying veneers of Sydney Basin outliers at Aarons Pass, Mt Margaret, the Castle
and Mt Bocoble (Fig. 12). At these localities, polymictic conglomerate and pebbly
litharenite represent the basal unit in the Snapper Point Formation of the Shoalhaven
Group (Bembrick, 1983).

Geological History of the Cudgegong-Mudgee District
Ordovician

The oldest rocks in the Cudgegong-Mudgee district, the Late Ordovician (Gis-
bornian) Cudgegong Volcanics consist of basaltic and andesitic arenite, breccia and rare
lava associated with volcaniclastic debris flows on the flanks of submarine volcanoes
with fringing shallow marine areas. These rocks are correlated with the upper parts of
the Sofala Volcanics in the Sofala district. Other strata tentatively assigned an Ordo-
vician age on the northern Capertee High are flysch-like sequences to the northwest
(Lue Beds, Offenberg ¢ al., 1971) and west of Rylstone (Colquhoun, 1989). The

PROC. LINN. SOC. N.S.W., 111 (3), 1989

196 CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

Cudgegong Volcanics-Sofala Volcanics association constitutes a Late Ordovician
volcanic arc providing a number of basaltic and andesitic eruptive centres with fringing
shallow water environments, with possibly deeper water flysch conditions to the east.

Silurian

The Cudgegong Volcanics are unconformably overlain by Wenlockian to Lud-
lovian sequences. Similar contacts occur at Sofala and possibly west of Rylstone, and the
absence of Llandoverian units confirms that deformation of the volcanic arc continued
through the Benambran/Quidongan event (Crook e¢ al., 1973; Cas, 1983).

There is a thick and persistent succession of Wenlockian to Ludlovian shallow
marine to possibly emergent units exposed in the Cudgegong-Mudgee district. The
lowermost unit, the Willow Glen Formation (conglomerate, pebbly arenite and arenite;
fossiliferous shale and limestone) was deposited in a southeast-facing coastal environ-
ment which included a fluvial channel zone; subtidal to supratidal flats, affected by a
series of transgressive-regressive cycles including common incision by fluvial channels;
and more open marine shelf conditions.

The conformably overlying Windamere Volcanics-Ioolamanang Formation
eruptive episode produced up to 1500m of undifferentiated dacite lava and breccia (the
Windamere Volcanics) possibly emplaced as a thick lava dome; and between 2 and 3km
of fine- to coarse-grained dacitic detritus with common fragmental basaltic blocks (the
Toolamanang Formation). The latter unit was produced by dense, gravity-driven
volcaniclastic ash- and mud-flows which incorporated eroded basement material. This
was followed by a period of volcanic quiescence represented by the Millsville Formation
(up to 220m of dominantly limestone with dacitic and rhyolitic detritus) deposited on a
shallow marine shelf.

Late Silurian shallow marine sediments with minor dacite/rhyolite lava and
fragmental rocks also occur to the north of Mudgee (Armstrong, 1983), west of Rylstone
(Colquhoun, 1989) and tentatively northwest of Rylstone (Offenberg e¢ al., 1971). In the
Sofala district, equivalents of the Willow Glen Formation (the Tanwarra Shale) and the
Windamere Volcanics-loolamanang Formation (the Bells Creek Volcanics) are known
(Packham, 1968); however, they are far more restricted in extent and thickness than on
the northern Capertee High. There is no Millsville Formation equivalent in the Sofala
district as the shallow marine rocks are conformably overlain by the deep water Hill End
Trough sequence.

Post-Silurian

Contacts with the Early Devonian strata are either faulted or disconformable, the
latter representing the effects of the Bowning deformational event (Cas, 1983) on the
northern Capertee High.

During the Early and part of the Middle Devonian, the deposition of shallow water
to emergent, richly fossiliferous sediments with silicic volcanic outpourings continued
on the northern Capertee High (Wright, 1966, 1967, 1981); however, the Queens Pinch
Belt consists of both shallow and deep water strata at the western margin of the Capertee
High, adjacent to the deep water Hill End Trough. The marked facies change to Late
Devonian fluviatile conditions is preceded by deformation resulting in the Lambian
Unconformity. Fluviatile conditions, with an increasing shallow marine component,
continued possibly into the Early Carboniferous when the effects of the cratonizing
Kanimblan deformational event (Cas, 1983) produced folding of the Late Devonian and
older rocks, and emplacement of the Middle Carboniferous Aarons Pass Granite.

Peneplanation of much of the Cudgegong-Mudgee district took place before the

PROC. LINN. SOG. N.S.W., 111 (3), 1989

J. W. PEMBERTON 197

Early Permian where the thin, flat-lying remnants of the basal Snapper Point Formation
of the Shoalhaven Group represent a sandy transgressive shoreline, at the western
margin of the Sydney Basin.

ACKNOWLEDGEMENTS

I gratefully acknowledge the continuing advice and support of Dr A. J. Wright
throughout the study. Dr Wright and Dr C. L. Fergusson kindly reviewed and construc-
tively criticized the manuscript. I thank all of the Department of Geology staff, as well as
Dr R. Offler, University of Newcastle, for their valuable and fruitful discussion. In
addition, the assistance and co-operation of various members of the Water Resources
Commission at Windamere Dam is acknowledged.

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WRIGHT, A. J., 1966. — Studies in the Devonian of the Mudgee district, N.S.W. Sydney: University of
Sydney, Ph.D. thesis, unpubl.

——,, 1967. — Devonian of the Capertee Geanticline, New South Wales, Australia. Jn OSWALD, D. H.., (ed.),
International Symposium on the Devonian system, Calgary. 1967. Alberta Soc. Petrol. Geol. 2: 117-121.

——,, 1981. — A new phillipsastraeinid tetracoral from the Devonian of New South Wales. Palaeontology 24:
589-608.

PROC. LINN. SOC. N.S.W., 111 (3), 1989

J. W. PEMBERTON

APPENDIX

199

Faunal Lists for the Willow Glen Formation

Locality Details

Locality 1
GR 618725; known locally as
Woolleys Flat

Locality 2
GR 642692 to GR 648687; area just
west of Ironstone Creek

Locality 3

GR 563740; just south and west of
four wheel drive track north of Lime-
stone Creck

Author (Source)

Pickett (1982a)

Strusz (pers. comm., 1984)

Pickett (1982a)

Strusz (pers. comm., 1984)

Pickett (1982a)

Strusz (pers. comm., 1984)

Fauna Recorded and Age

corals: Syringopora, Halysites ortho-
pteroides, Desmidopora multitabulata,
Pycnostylus scalariformis
brachiopods: pentamerids
conodonts: Panderodus, Ozarkodina
ranuliformis, Distomodus, ‘Ozarkodina’
media, ‘Neoprioniodus’ cf. bicurvatus
age: Wenlockian to Ludlovian

brachiopods: Spirinella, Aegiria cf.
norvegica, ?Howellella, Coelospira, Salo-
pina, Stropheodontacea

trilobites: Encrinurus mitchellz,
proetacean, ?Staurocephalus struszi
(cheek only)

crinoids: Pisocrinus

age: Late Wenlockian to Early
Ludlovian

corals: Favosites, Heliolites, Tryplasma,
Alveolites, Thamnopora
stromatoporoids: Amphipora
brachiopods: Kirkidium

conodonts: Ligonodina} “?Lonchodina;}
Panderodus, Ozarkodina ranuliformis
age: Wenlockian to Ludlovian

brachiopods: Spzrinella, ?Howellella,
Coelospira cavata, Salopina ?medtocostata,
Morinorhyncus oepiki, Maoristrophia,
Leptostrophia

trilobites: Encrinurus mitchellz,
proetacean

corals: Entelophyllum

age: Late Wenlockian to Early
Ludlovian

corals: Phaulactis, Tryplasma, Alveolites
stromatoporoids: Amphipora
brachiopods: Kzirkidium

conodonts: Ozarkodina ranuliformis
age: Wenlockian to Ludlovian

brachiopods: Aegzria, Coelospira,
?Howellella

trilobites: Encrinurus mitchelli
corals: Halysites, Tryplasma
age: Late Wenlockian to Early
Ludlovian

PROC. LINN. SOC. N.S.W., 111 (3), 1989

200

Locality Details

Locality 4
GR 600732; just west of Horseshoe
Bend on Cudgegong River

Locality 5

GR 653637; in the upper limestone
bed just west of track along Oakey
Creek, southeast of Cudgegong

Locality 6
GR 560761; approx. 2km west of
‘Windamere Village’

Locality 7

GR 612698; just north of new
Cudgegong-Mudgee road, approx.
3km east of Limestone Creck

Locality 8

GR 562717; approx. ikm south of
Limestone Creck road, at cast margin
of southwest areca

Locality 9
GR 528796; approx. 0.5km southeast
of Appletree Flat, northwest area

PROG. LINN. SOG. N.S.W., 111 (3), 1989

Author (Source)

Strusz (pers.comm., 1984)

Pemberton (additional
specimens identified by
A. J. Wright)

Pickett (1982a)

Pemberton (identified
by A. J. Wright)

Pemberton (identified by
A. J. Wright)

Pickett (1982a)

Pemberton (identified by
A. J. Wright)

CUDGEGONG-MUDGEE DISTRICT STRATIGRAPHY

Fauna Recorded and Age

corals: Cystaphyllum, Tryplasma
?lonsdaler, Phaulactis shearsby,
Syringopora, Palaeophyllum

corals: Halysites, Favosites, Heliolites
age: probably Wenlockian to
Ludlovian

corals: Propora, Tryplasma, Favosites,
Phaulactis

brachiopods: Kzrkidium

age: Wenlockian to Ludlovian

corals: Tryplasma, Thamnopora,
Phaulactis, Favosites, Halysites
brachiopods: pentamerids
age: Late Silurian

corals: Halysites, Favosites
brachiopods: pentamerids
age: Late Silurian

conodonts: ‘Ligonodina; Spatho-
gnathodus; ‘Hindeodella; ‘Ozarkodina’
denckmanni, Distomodus, Panderodus,
Ozarkodina cf. remschetdensis, Delotaxts
cf. elegans, ‘Belodella’ triangularis
age: very Early Devonian

corals: Halysites, Favosites, Helvolites
brachiopods: pentamerids

age: Late Silurian

corals: Helzolites
brachiopods: pentamerids
age: Late Silurian

Aspects of the History and Fishery of the
Murray Cod, Maccullochella peels
(Mitchell) (Percichthyidae)

STUART J. ROWLAND
(Communicated by J. R. MERRICK)

ROWLAND, S. J. Aspects of the history and fishery of the Murray cod, Maccullochella peeli
(Mitchell) (Percichthyidae). Proc. Linn. Soc. N.S.W. 111 (3), 1989: 201-213.

Aspects of the history and fishery of Australia’s most famous and largest inland
freshwater fish, the Murray cod, Maccullochella peeli, are briefly reviewed. Information
and data on fossil records, the prominence of Murray cod in aboriginal mythology and
culture, observations of cod by explorers and early settlers, and the development and
subsequent decline of a commercial fishery are presented. Possible factors contributing
to the reduced abundance of Murray cod are discussed. It is suggested that overfishing
caused a decline between the late 1800’s and the 1930’s, but that extensive environmen-
tal modification of the Murray-Darling river system has adversely affected larval
recruitment resulting in the dramatic decline in abundance of M. peel since the 1950’s.

Stuart J. Rowland, NSW Agriculture G Fisheries, Eastern Freshwater Fish Research Hatchery,
Agricultural Research & Advisory Station, Grafton, Australia, 2460; manuscript received 20 July
1988, accepted for publication 19 April 1989.

INTRODUCTION

The Murray cod, Maccullochella peeli (Mitchell, 1838) is an Australian, native,
warmwater, percichthyid fish found naturally throughout most of the Murray-Darling
river system (Fig. 1) with the exception of the headwaters of some tributaries in Victoria
and southern New South Wales (Lake, 1971).

M. peeli is Australia’s, and one of the world’s largest freshwater fish. Whitley (1955)
stated that Murray cod grow to 1.8 m (6 ft) and 83 kg (182 lb); however, a cod of 113.6 kg
(250 lb) is reported to have been captured from the Barwon River near Walgett in 1902
(Noble, 1955). Although cod in excess of 50 kg are rarely captured, small numbers of cod
between 20 and 40 kg are regularly taken by experienced commercial and recreational
fishermen, particularly in the Darling, Barwon and Edward rivers, the lower reaches of
the Murray and Murrumbidgee rivers, Lake Mulwala on the Murray River, and Lake
Burrinjuck on the headwaters of the Murrumbidgee River. Because of its size and
excellent edible qualities, Murray cod is highly valued by both commercial and recre-
ational fishermen.

Despite its importance, there has been little research into the natural history of MZ.
peelt. This paper briefly reviews the part played by Murray cod in aboriginal mythology
and culture, the observations made of the species by explorers and early settlers, and the
development, decline and current status of the Murray cod fisheries. Possible causes of
the dramatic decline in abundance of Murray cod are discussed.

HISTORICAL ASPECTS OF MURRAY COD

Ongin and Fossil Records

Most Australian freshwater fishes, including M. peeli, are considered to have a
relatively recent marine ancestry (Whitley, 1959; Darlington, 1965). MacDonald (1978)
suggested that the Maccullochella and Macquaria groups (both percichthyids) diverged
from a common ancestor during a marine stage of their evolutionary development and
made separate colonizations of Australian freshwaters.

Hills (1946) recorded fossil Murray cod from diatomaceous earth in the Warrum-

PROC. LINN. SOC. N.S.W., 111 (3), 1989

202 MURRAY COD HISTORY AND FISHERY

bungle Mountain areas and considered the remains to be not older than Pliocene
(started 7 m.y.a.). However, Browne (1972) stated that the basalt overlaying these dia-
tomaceous earths had been dated as Upper Miocene, and Taylor et al. (1980) reported
Murray cod fossils from diatomite in the Cooma region to be of Miocene age (26-7
m.y.a.). The diatom flora associated with these fossils is of the type found with Lower
Tertiary basalts (Gill, 1970) and so the ancestral Maccullochella may be up to 60-65
million years old. Hills (1946) stated that the former distribution of MZ. peel (M.
macquariensis as used by Hills, 1946, synonymous with M. peelz) in the western drainage,
as determined by fossil records, was similar to the current distribution.

aie

MURRAY- | Condamine R

DARLING,

SYSTEM “\y Amn) ff 7 BRISBANE

Warrego R

3 ]

A i

rf
i

| ae

ne r

LC
:

Fig. 1. The Murray-Darling river system.

Aboriginal Mythology and Culture

The Murray cod plays a prominent part in aboriginal mythology (Ramsay Smith,
1930; Berndt, 1940). According to legend, a huge fish, the Murray cod (called ‘ponde’ by
the aborigines*) burst forth from the depths of the earth at the source of the Murray

* According to Ramsay Smith (1930) and Berndt (1940) the aborigines of the lower Murrumbidgee and
Murray River regions called the Murray cod ‘ponde’, but Bennett (1834) states that aborigines in the Yass
region on the upper Murrumbidgee River called the river cod ‘mewuruk’ and the aborigines in the Tumat [sic]
country called the varieties of river cod ‘bewuk’ or ‘mungee’.

PROC. LINN. SOC. N.SW., 111 (3), 1989

S. J. ROWLAND 203

River, which was then only a small stream of water trickling to the southern ocean. The
Murray cod struggled along the narrow stream digging with its head and swinging its
powerful tail making the river deep and forming all the bends. Then Nepelle, the Great
Prophet, speared it at a’site now known as Lake Alexandrina and with the help of the
creative hero Ngurunderi, cut it into pieces and threw the fragments into the water,
naming them ‘tarki (golden perch, Macquaria ambigua), ‘tukkeri (bony bream, Nematolosa
erebr), ‘tinuwarre (silver perch, Bzdyanus bidyanus) and all the other fishes of the system.
When they had finished they threw the remainder back and said “You keep on being
ponde’.

Murray cod were a major food item of those tribes living adjacent to inland waters
(Lawrence, 1971; Tindale, 1981) and as with all animal life, the aborigines made a
detailed study of inland fishes, in particular the Murray cod which was considered the
fish (Ramsay Smith, 1930). The aborigines were excellent fishermen and amazed early
explorers and settlers with their prowess. They commonly used spears, nets and poisons
to capture their prey, but also constructed different types of traps using brush fences,
stones or hollow logs (Bennett, 1834; Lawrence, 1971; Tindale, 1951, 1981).

Explorers and Early Settlers

The inland explorers and early settlers were astounded by the abundance, size and
delicacy of the Murray cod or as it was generally known by the pioneers ‘River cod’ or
‘codfish’. The explorer John Oxley (1820) wrote of cod in the Lachlan River ‘If however
the country itself is poor, the river is rich in the most excellent fish, procurable in the
utmost abundance. One man in less than an hour caught eighteen large fish, one of
which was a curiosity from its immense size and the beauty of its colours. . . It weighed
an entire 70 pounds, . . . Most of the other fish taken this evening weighed from fifteen
to thirty pounds each’.

The holotype of M. peelz, which has since been lost (Berra and Weatherley, 1972),
was collected from the Peel River, N.S.W., by the explorer Major Thomas Mitchell
(Mitchell, 1838). There is an excellent drawing, dated 14th December, 1831, of a Murray
cod in Major Mitchell’s sketch book. Members of the expedition led by Charles Sturt
down the Murrumbidgee and Murray rivers, caught and ate Murray cod (Sturt, 1899).

Bennett (1834) wrote that large quantities of the delicious ‘River Cod’ weighing up
to 120 lbs were caught in the Yas [sic] and Murrumbidgee rivers, and in 1863 he recom-
mended to the Acclimatization Society of N.S.W. that every endeavour should be made
to propagate them (Bennett, 1864). Murray cod were held in such high esteem that Dr
Gunther considered the species worthy of acclimatization in England (O’Connor, 1897)
and Ramel (1868; cited in Berra and Weatherley, 1972) suggested that Murray cod be
introduced into Europe. Although this did not eventuate, Murray cod were stocked,
during the 19th century into many waters where they were not found naturally. These
included the Yarra River, Victoria (Wilson, 1857), the easterly-flowing Cox’s, Nepean
and Wollondilly rivers on the N.S.W. central coast (Phillips, 1863; Hill, 1864), Mul-
warree Ponds near Goulburn and Lake George near Canberra (Macleay e¢ al., 1880), the
easterly-flowing Mary river system in southern Queensland (S. H. Midgley, pers.
comm.), and the Avon River and Lake Grassmere, Western Australia (Morrissy, 1970).
Apart from an occasional specimen captured from the upper reaches of the Yarra River,
Murray cod are no longer found at any of these sites. The population of cod in the Mary
river system is genetically and morphometrically distinct from M. peeli, but its
taxonomic status has not been determined (Rowland, 1985).

During the 20th century, Murray cod have been successfully stocked into Lake
Bathurst and Lake George in N.S.W. (Whitley, 1937), Lake Charlegrark, Green Lake,
Taylor’s Lake and the Wimmera River, in the western Wimmera region of Victoria

PROC. LINN. SOC. N.S.W., 111 (3), 1989

‘TI6] ‘YuvuuoY rwou sary Aesunpy oy} UO AOMOTARAY, JoUIv9}So[pped oy ps

voqde YIIBI OUT, Buy

Vii

MURRAY COD HISTORY AND FISHERY

204

PROC. LINN. SOG. N.S.W,, 111 (3), 198

S. J. ROWLAND 205

(Cadwallader and Backhouse, 1983), Cataract Dam (Anderson, 1916) and several other
Sydney water supply dams (Lake, 1959, 1971) and numerous farm dams in the eastern
states of Australia.

MURRAY COD FISHERIES

Development

During the mid to late 1800’s a large inland, commercial fishery developed and was
based mainly on the Murray and Murrumbidgee rivers (Macleay et al., 1880; Dannevig,
1903; Stead, 1903). Large-scale operators used paddlesteamers as fishing boats (Fig. 2)
and set up to 200 drum nets per vessel (Pollard and Scott, 1966). The drum net was
introduced into the inland fishery in 1880 (Dannevig, 1903) and has remained the most
common method of netting Murray cod and golden perch in the inland rivers and
creeks. By 1883 the Murray River fisheries formed a considerable factor in the fish
supply to Victoria and during this year more than 147 tons were sent to Melbourne from
Moama (Cox, 1884). In 1900 the value of the inland fisheries within the South
Australian portion of the Murray River was worth £25,000-£30,000 per annum
(Whitley, 1937).

Although few quantitative data are available, it appears that the fishery was based
primarily on Murray cod. In 1862 a company of six men and a number of aborigines
captured two to three tons of fish per week from the Murray River, south of Deniliquin,
which ‘abounded with fish, particularly the Murray River cod’ and sent them to Bendigo
and Melbourne (Jervis, 1952). A professional fisherman from Tailem Bend, South
Australia, reported catching’. . . three bags (170 lb each) of Murray cod in one morning’
(Stead, 1903). The Fisheries Enquiry Commission of 1879-80 (Macleay et a/., 1880) was
told by Mr. F. A. Tompson of Wagga, in relation to the fish in the Murrumbidgee River,

that “The cod is the most prominent and remarkable . . . It is brought to market more
plentifully than the others . . . a ton of fish is brought in here every week . . . I saw 150
large cod alive inacart. . . They were sold in two hours or less’.

In 1900, cod accounted for 75% of the river fish available at the Melbourne market,
the remainder being golden perch (Poole, 1984). The dominance of Murray cod in
professional catches is shown in Fig. 2, the catch aboard the paddlesteamer ‘Mayflower’
near Renmark in 1911. Dakin and Kesteven (1938) referred to ‘the overwhelming impor-
tance of cod, the quantity of golden perch and silver perch taken is small compared with
quantity of cod’.

Besides the commercial fishery, large numbers of Murray cod were easily caught
and used as food by early settlers (Bennett, 1834; Macleay et al., 1880). By 1955 the
popularity of inland fishing had increased tremendously and with the aid of motor
vehicles, a large recreational fishery had developed (Anon., 1956). During the 1950’s
large catches of Murray cod could be made by experienced recreational fishermen.

Decline

Dakin and Kesteven (1938) presented the available data on the catch of native fish
from the inland waters between 1883 and 1938, and allowing for the limitations as
discussed by those authors, the data indicate that although large fluctuations of fish
populations occurred in the Murray-Darling river system, there had been a gradual
decline in the overall catch from a peak in 1918. By the mid 1930's it was apparent that
the commercial fishery had declined to an unprofitable level for the large-scale operators
(Whitley, 1937; Pollard and Scott, 1966).

Catch statistics from the commercial fishery in inland N.S.W. between 1940/41 and
1983/84 are presented in Fig. 3. The annual catch of Murray cod increased between

PROC. LINN. SOC. N.S.W., 111 (3), 1989

206 MURRAY COD HISTORY AND FISHERY

(a) 150
co} 100
fo)
3)
>
oO
4
Goa
30
= x
wo
23 50
ad
)
yp x
ow
iS)
(b)
1000

~
uo
Oo

per licenced boat
(kg)
N uo
uo (=)
oO fo)

Catch of Murray cod per
licenced fisherman and

c) :
| 100
a

2 75
°

4 : 50
o

c oa

o 5

oO Ta

i 25
Ay

1940/41 1950/51 1960/61 1970/71 1980/81

Fig. 3. The total annual catch (a) and catch per licenced fisherman and per licenced boat (b) of Murray cod,
and the percentage composition of Murray cod, golden perch and common carp (c) in the commercial catch
from inland waters in N.S.W. between 1940/41 and 1983/84 (data from annual reports, N.S.W. State

Fisheries).
(b) O © catch per licenced fisherman (c) M——W@ = Murray cod
A ------ A catch per licenced boat (J——U _-s golden perch

=> common carp

PROG. LINN. SGC. N.S.W., 111 (3), 1989

S. J. ROWLAND 207

1940/41 and 1955/56, but then the total catch and the catch per licenced fisherman
declined dramatically (Fig. 3a, b). Murray cod was the major species in the inland
fishery until 1951/52, and between 1940 and 1951 cod comprised 42-65% of the total
annual catch from inland N.S.W. However, after 1951/52, golden perch (Macquaria
ambigua) replaced Murray cod as the major native species in the N.S.W. inland fishery
(Fig. 3c). The total catch, catch per licenced fisherman and the percentage composition
data all strongly suggest that there was a dramatic decline in the abundance of M. peeli
between 1955 and 1964. There was a concurrent decline in the commercial catch of
Murray cod from the Murray River in South Australia (Reynolds, 1976). Between
1940/41 and 1962/63 the annual total catch of Murray cod in N.S.W. exceeded 35,000 kg;
however since 1963/64, with the exception of 1974/75, the annual catch has remained
below 30,000 kg and the catch per licenced fisherman below 200 kg.

The continued small catch and low catch per licenced fisherman of Murray cod in
the N.S.W. commercial fishery since 1960/61 (Fig. 3) indicate that the fishery remains in
a depressed state. These catch statistics also indicate that there has been no major
reduction or increase in the abundance of Murray cod since the mid 1960's.
Consequently, it appears that stocks of Murray cod have remained stable, but at
relatively low levels over the last 25 years.

Reduced Distribution

There has also been a reduction in the distribution of M. peels. Many historical
reports indicate that the species was common in rivers where cod are now rare or no
longer found. The explorer George Evans, the first European man to see the species
(Stanbury and Phipps, 1980), observed cod in the Fish River before he reached the
present site of Bathurst. The type locality of M. peeli is the Peel River, N.S.W. probably
near the present site of Tamworth (Mitchell, 1838) and in 1836 an enormous 120 lb
‘River codfish of the Colonists’ was found entangled and struggling near the bank in a
pond of the Cudegong River (Bennett, 1864). Murray cod are now extremely rare in the
Fish, Peel and Cudegong rivers.

In Victoria, Murray cod abounded in the Loddon, Campaspe and Goulburn rivers
and their tributaries, even where the waters ‘dwindled into the most insignificant
streams’ (Wilson, 1857). By the late 1940’s the populations of Murray cod and other
native fishes had declined in these rivers (Langtry, in Cadwallader, 1977) and there are
now very few localities in Victoria where Murray cod can be considered common
(Cadwallader and Backhouse, 1983).

Fisheries Management and Research

Concern about the stocks of Murray cod was expressed as early as the 1880 Royal
Commission enquiring into the Fisheries of N.S.W. (Macleay et al., 1880) and from 1883
until about 1895 there was some supervision of inland waters (Dakin and Kesteven,
1938). Dannevig (1903) detailed measures for restricting gear, protecting fry and young
fish and the imposition of a closed season. From 1905 to 1910 serious attempts were
made to improve the cod fishery, including experiments conducted in 1905 by H. C.
Dannevig on the artificial propagation of cod (Farnell, 1906; Dakin and Kesteven,
1938). Whitley (1937) briefly discussed the available information on the distribution,
fishery, breeding and taxonomy of Murray cod and included a complete bibliography
containing mostly taxonomic references. In 1936, a conference on the Murray River
fisheries, attended by representatives from N.S.W., Victoria and South Australia,
adopted a closed season of September, October and November for the taking of Murray
cod; set minimum legal lengths; suggested that hatcheries be established; asked the
Murray River Commission to construct a fishway at Lock 15 at Euston; and suggested

PROG. LINN. SOG. N.S.W,, 111 (3), 1989

208 MURRAY COD HISTORY AND FISHERY

that the breeding habits and migration of freshwater indigenous fishes be studied by
each State (Isherwood, 1939).

However, since that time only five studies have contributed significantly to the
knowledge of the biology of M. peeli. Dakin and Kesteven (1938) presented brief notes on
the natural history, behaviour in captivity and the spawning season of Murray cod; they
also discussed the cod fishery including the evidence for, and possible causes of the
decline of cod stocks. Dakin and Kesteven artifically bred cod by capturing and strip-
ping ripe fish; the eggs, embryonic development, larvae and fry were described.
However, this part of the study was restricted by the difficulty of procuring ripe brood-
fish from the wild. The need for future research into the biology and breeding of this
species was emphasized.

J. O. Langtry conducted an ecological survey of the Murray River and some of its
tributaries in 1949-50. Unfortunately, his report was not published, and it wasn’t until
1977 that the manuscript was rewritten and presented by Cadwallader (1977). The
report contains data on the relative abundance of fishes in the study area and describes
the differences between Murray cod and trout cod (Maccullochella macquariensis). Langtry
also made observations and collected some quantitative data on the distribution, diet,
breeding biology and growth rate of Murray cod.

During the 1960’s, John Lake studied the reproductive biology of native fishes at
the Inland Fisheries Research Station, Narrandera, N.S.W. and his research demon-
strated that critical temperatures and rising water levels in ponds (and presumably a
flood or fresh in the wild) triggered the spawning of some species (Lake, 1967 a,b).
Cadwallader and Gooley (1985) collected data on the spawning of Murray cod in
earthen ponds, and developed techniques for the artificial propagation and rearing of
M. peeli. Rowland (1985) conducted research into the biology and artificial breeding of
M. peeli, and some of his findings form the basis of the following discussion.

POSSIBLE CAUSES OF THE DECLINE

Reduced Larval Recruitment

The high survival of fish larvae is dependent on the availability of relatively high
concentrations of suitable-sized food organisms at the commencement of exogenous
feeding; suboptimal feeding conditions generally result in death due to starvation or
predation (May, 1974; Pitcher and Hart, 1982). Hjort (1926) hypothesized that the
degree of mortality of larvae during a ‘critical period’ after the completion of yolk sac
absorption determined the strength of year-classes in natural populations. Although the
relationship between larval mortality during the ‘critical period’ and year-class strength
is difficult to determine in nature (May, 1974) it is generally thought that the survival
rate of fish larvae is the most important factor determining the strength of year-classes
(Beverton, 1962; Gulland, 1965; May, 1974; Pitcher and Hart, 1982).

Many overseas studies have shown that strong year-classes of freshwater fishes are
established when the breeding season coincides with rising or high water levels (e.g.
Agegus and Elliott, 1975; Stevens, 1977; Marshall, 1982; Beam, 1983). Although M. peel:
spawned annually in the southern tributaries of the Murray-Darling river system
between 1977 and 1980, relatively strong year-classes were only established when the
breeding seasons coincided with high river levels or floods (Rowland, 1985), demon-
strating that floods in October and November provide optimum conditions for the
survival and recruitment of M. peeli larvae.

The floodplain areas of the Murray-Darling river system are highly productive.
When they are inundated in spring or summer, a rich source of terrestrial nutrients,
plus the plankton, aquatic insects (in particular chironomid larvae) and other organisms

PROC. LINN. SOG. N.S.W., 111 (3), 1989

S. J. ROWLAND 209

of the billabongs become available to the aquatic community of the rivers (Frith, 1959;
Shiel, 1980; Maher and Carpenter, 1984). Zooplankton, chironomid larvae and other
aquatic insects are the major food items of the larvae and fry of Murray cod and golden
perch in earthen ponds, and a delay of several days in the availability of food to larvae
after the completion of yolk sac absorption results in reduced survival in both species
(Rowland, 1985, 1986).

The construction of dams, high-level weirs and levee banks on the major tributaries
of the Murray-Darling system has altered the natural flow and temperature regimes and
dramatically reduced the frequency, extent and duration of floods (Lake, i971;
Reynolds, 1976; Cadwallader, 1978; Walker et al., 1978; Walker, 1979). The Murray
River no longer floods annually in spring and much of the vast anabranch, billabong
and floodplain areas of the Murray and Murrumbidgee rivers have been eliminated and
do not flood except under extraordinary circumstances (Langtry, in Cadwallader, 1977;
Shiel, 1980). Consequently optimum conditions for the survival of the larvae of Murray
cod, which usually spawn in October or November, now rarely occur. It is suggested that
the reduced frequency, extent and duration of spring flooding in the Murray-Darling
river system has led to an overall reduction in larval recruitment and that this is a major
cause of the decline in the abundance of M. peelz.

This hypothesis is supported by the change in the relative proportion (by weight) of
Murray cod and golden perch in the commercial catch from inland N.S.W. (Fig. 3c).
Prior to 1951/52, M. peel: was the major species; however, since then golden perch have
formed a much greater percentage of the annual catch than have Murray cod. Golden
perch require a substantial rise in water level, when temperatures are about 23°C, to
induce spawning (Lake, 1967a); if suitable conditions do not occur in the wild, adults
remain at an advanced stage of gonadal development until March or April (Mackay,
1973). Therefore M. ambigua can delay spawning over a six month period whereas M.
peelt spawns only during spring and early summer when the water temperature is about
20°C (Rowland, 1983, 1985; Cadwallader and Gooley, 1985). The reduced frequency
and extent of flooding, in particular the reduction of flooding which usually occurred
each spring in the Murray and Murrumbidgee rivers (Langtry, in Cadwallader, 1977;
Walker e¢ al., 1978; Walker, 1983) would be expected to affect larval recruitment to a
greater extent in M. peel: than in M. ambigua.

The major impoundments on the Murray-Darling river system in N.S.W. were
constructed between 1907 and 1976; Burrinjuck Dam 1907 (enlarged 1957), Hume Dam
1936 (enlarged 1961), Wyangla Dam 1936 (enlarged 1971), Yarrawonga Weir 1939,
Keepit Dam 1960, Menindee Lake Storage Scheme 1960, Burrendong Dam 1967,
Blowering Dam 1968, Pindari Dam 1969, Copeton Dam 1976 (Anon., undated; Walker,
1980). By 1950, the effects of Hume Dam and Yarrawonga Weir on the flow in the
Murray River were clearly evident (Langtry, in Cadwallader, 1977). The cumulative
effects of the major impoundments and water storage schemes would have been
substantial by 1960 when there was an apparent rapid decline in the abundance of
Murray cod.

Overfishing

The relatively large inland commercial fishery which existed between the mid
1800’s and the late 1930’s and which was based primarily on Murray cod (Macleay et al.,
1880; Dannevig, 1903; Stead, 1903; Dakin and Kesteven, 1938) would have placed
intense fishing pressure on cod populations. Previously unfished populations of long-
lived fishes consisting of 12 or more year-classes are extremely susceptible to exploi-
tation; and with unchanging recruitment the absolute size of the total stock will decline
markedly, or even catastrophically under moderate exploitation (Ricker, 1963). It is

PROG. LINN. SOG. N.S.W., 111 (3), 1989

210 MURRAY COD HISTORY AND FISHERY

therefore probable that the decline in abundance of Murray cod, at least until the 1930's,
by which time the inland commercial fishery had become unprofitable for large-scale
operators (Pollard and Scott, 1966), was caused primarily by overfishing.

A possible reduction in the exploitation of cod populations after the 1930’s due to
the depressed state of the fishery, the Depression and World War II, may have resulted in
numbers and the catch per licenced fisherman/boat remaining reasonably stable or even
increasing (Fig. 3) until the late 1950’s when, as suggested, the effects of reduced larval
recruitment became apparent by the rapidly declining stocks. It is also possible that the
recreational fishery which developed in the 1950’s (Anon., 1956; Poole, 1984)
contributed to the decline of cod stocks during this period.

English Perch (Redfin)

English perch, Perca fluviatilis, were abundant and sympatric with M. peel: in the
southern tributaries of the Murray-Darling system between the late 1940’s and the
1960’s. Langtry (in Cadwallader, 1977) found that the diet of P fluviatilis was identical to
that of M. peeli and M. ambigua, and small fishes become a major part of the diet of larger
English perch (Lake, 1967c). English perch larvae and juveniles feed on zooplankton,
crustaceans and insect larvae, and because P. fluviatilis spawns in early spring when
temperatures are about 12°C; usually the last week or so in August in southern N.S.W.
(Lake, 1967c), juvenile English perch may prey on, and possibly compete for food with
Murray cod larvae and fry, particularly during drought periods when food resources are
limited. The catch data on fishes from the Kerang Lakes, Victoria, between 1919 and
1949 (Figure 1 in Cadwallader, 1977) demonstrated that when English perch were
abundant, native fish were scarce and vice versa. It is therefore possible that English
perch have contributed to the decline of Murray cod in the southern parts of the
Murray-Darling river system.

Other Factors

Factors such as siltation, desnagging, channelization, depressed water tempera-
tures below impoundments, and barriers preventing spawning migrations have been
implicated in the decline of native fishes (see Merrick and Schmida, 1984). While some
of these factors may be responsible for the decline of some species in certain areas,
because of the great size of the Murray-Darling river system it is unlikely that they have
been major contributing factors to the decline of M. peels which is widely distributed

throughout the system (Lake, 1971) and does not undergo extensive migrations
(Reynolds, 1983).

SUMMARY

Historical aspects of Australia’s most famous and largest inland freshwater fish, the
Murray cod, Maccullochella peeli, are briefly reviewed. Fossil records suggest that
ancestral Murray cod are of, at least Miocene age. The Murray cod played a prominent
part in the mythology and culture of some aboriginal tribes. Explorers, including
George Evans, John Oxley, Thomas Mitchell, and Charles Sturt, and the early, inland
settlers were astounded by the abundance, size and delicacy of Murray cod. The species
was held in such high esteem that it was considered worthy of acclimatization in
England and Europe, and although this did not eventuate M. peel: was extensively trans-
located within and outside its natural range of the Murray-Darling river system during
both the 19th and 20th centuries.

During the late 1800's a large, inland, commercial fishery developed. It was based
on Murray cod and was located mainly on the mid and lower reaches of the Murray and

PROC. LINN. SOC. N.S.W., 111 (3), 1989

S. J. ROWLAND 211

Murrumbidgee rivers. Historical records and catch statistics indicate that there has
been a dramatic decline in the abundance and a reduction in the distribution of M. peelz.
Possible causes of the decline are briefly discussed. It is suggested that overfishing
contributed to a decline between the late 1800’s and the 1930's, but that the reduced
frequency, extent and duration of spring flooding in the Murray-Darling river system,
caused by the construction of dams, high-level weirs and levee banks, has adversely
affected larval recruitment in M. peeli. This has caused the dramatic decline in
abundance of cod during the 1950’s and the maintenance of stocks at relatively low levels
over the last 25 years.

ACKNOWLEDGEMENTS

I gratefully acknowledge Ray Gregg for providing the photograph of the paddle-
steamer ‘Mayflower’. I express my appreciation to Peter Williamson and David Rodgers
for preparing the figure and photograph respectively, and to Margaret Leedow and
Denis Reid for providing the fisheries catch statistics. I thank Barbara Butler for typing
the manuscript.

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PROC. LINN. SOC. N.S.W,, 111 (3), 1989

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Microhydromys musseri n. sp., a New Murid
(Mammalia) from the ‘Torricelli Mountains,
Papua New Guinea

T. F. FLANNERY

FLANNERY, T. F. Microhydromys musseri n. sp., a new murid (Mammalia) from the
Torricelli Mountains, Papua New Guinea. Proc. Linn. Soc. N.S.W. 111 (3), 1989:
215-222.

Microhydromys mussert n. sp. differs greatly from the only previously described
species of Microhydromys (M. richardsonz) in its larger size, brown dorsum with a markedly
contrasting venter, broader skull, palate morphology, and dentition details. Both
species however share synapomorphies not seen in other hydromyine murids. Micro-
hydromys mussert n. sp. is thus far known from a single specimen collected near the
summit of Mt Somoro, Torricelli Mountains (part of the North Coast Ranges). It is the
third mammal species endemic to these ranges to be described. Three additional speci-
mens of Microhydromys richardsoni, which was previously known only from the holotype,
are described. These add significantly to knowledge of the species distribution and
morphology.

T. F. Flannery, Australian Museum, Sydney, Australia 2000; manuscript received 31 January
1989, accepted for publication 19 April 1989.

KEY WORDS: Microhydromys mussert, New Guinea, Hydromyinae, zoogeography.

INTRODUCTION

The murid subfamily Hydromyinae consists of 10 genera in the Australo-Papuan
region. Thomas (1898) and Misonne (1969) have referred Asian taxa to the Hydro-
myinae, but Musser (1982) notes that it is not yet clear if any of these taxa do indeed
represent part of the hydromyine radiation. All of the Asian taxa are readily distinguish-
able from members of the Australo-Papuan group and none resemble the species of
Mucrohydromys. For this reason, and because of their uncertain status, I have restricted
comparisons of the new species to Australo- Papuan taxa.

Within the New Guinean Hydromyinae are a group of six species, placed in four
genera, that are known as microhydromyines. They are small, shrew-like murids with
short, sleek coats, tiny eyes and pointed snouts. Included are some of the rarest and least
known Melanesian mammals. The genera Mayermys and Neohydromys are monotypic,
while Pseudohydromys includes two similar, probably parapatric species. Microhydromys
includes two species that differ greatly in morphology, one of which is described here as
new.

Previous taxonomic work upon microhydromyines has been limited, and it is not
yet clear whether these species represent a monophyletic group, or are simply pheneti-
cally similar and paraphyletic. Likewise, the generic limits are in some cases poorly
understood, and it is possible that in future some genera will be synonymized.

Microhydromys richardsoni Yate and Archbold, 1941 was described on the basis of a
single adult male taken at an altitude of 850m near Bernhard Camp, Idenburg River,
Irian Jaya. It is one of the more distinctive microhydromyines, possessing grooved upper
incisors and a broad skull with a short rostrum. A brief mention of the existence of an
additional specimen (Menzies and Dennis, 1979) constitutes the only other reported
occurrence of this species.

In this work a new species of Microhydromys, from the North Coast Ranges, is
described; descriptions of 3 additional specimens of M. richardsoni are provided.
Although the new species is known only from a single specimen its description as a new

PROC. LINN. SOC. N.S.W., 111 (3), 1989

216 A NEW MURID FROM PAPUA NEW GUINEA

taxon is considered justified on the basis of its highly distinctive morphology. The likeli-
hood of additional material being found in the near future is also low.

MATERIALS AND METHODS

Dental terminology follows Musser (1981), and colours where capitalized follow
Smithe (1974). Measurements are in millimitres and weights in grams. Abbreviations
are as follows: AM M, Australian Museum mammal specimen; AMNH, American
Museum of Natural History mammal specimen; BBM, Bishop Museum mammal
specimen.

SYSTEMATICS
Maucrohydromys ‘Yate and Archbold, 1941
Maicrohydromys mussert n. sp

(Figs 1-2, Table 1)

Holotype and Type Locality: BBM101737, puppet skin and skull with dentaries of an
adult male, collected on 12 November 1972 by A. B. Mirza at 1,350m on Mt Somoro (3°
22'S, 142° 09’E), Torricelli Mountains, West Sepik Province, Papua New Guinea.

Etymology: For Dr Guy Musser, who has contributed so abundantly to systematic
research on the murids of Asia and Melanesia, and who has so greatly increased our
understanding of the shrew-like murids of the region.

Diagnosis: Microhydromys musseri n. sp. can be distinguished from M. richardson: in the
following ways: it is larger (Table 1); the dorsum is Ochre Brown and the sharply demar-
cated venter Cinnamon (as opposed to the nearly uniform grey colouration of M. richard-
soni); there is no longitudinal groove on the upper incisors; the palate is more deeply
concave; the skull is absolutely and relatively broader; I/1 is less procumbent; a posterior
cingulum is present on M/1.

Description: Fur short and dense, tail and ears appearing naked to the unaided eye.
Dorsum near Ochre Brown, grading to Cinnamon Brown on flanks. Venter sharply
demarcated and Cinnamon with two small, irregular white patches on thorax. Throat
Cinnamon, crown of head Fuscous. Eyes surrounded by a circle of blackish hairs. Most
vibrissae pale, but a few black; similar in length, distribution and density to those of
other microhydromyines such as M. richardsont and N. fuscus. Hands and feet thinly
furred with pale hairs on dorsal side. Tail slightly paler below than above and mottled
with light patches becoming prominent distally. Flesh of ears dark. Tail scales ill-defined
but form regular rings with a single hair per tail scale. Flesh of the hands and feet
shrivelled, but enough can be seen to determine that the pads of the forefeet were
striated. Tiny rhinarium naked, toes unwebbed.

Upper incisors ungrooved and enamel orange. Lower incisors with paler orange
enamel and less procumbent than in most other hydromyines. Molars heavily worn,
with much crown detail lost. M1/ elongate, narrow, subrectangular. T2 large, obvious,
much worn. A slight groove on the anterolingual face separates Tl and T2. Distinct
enamel basin separates anterior and median lophs. Wear has obliterated almost all
detail distal to this point. However, a slight groove discernable between median and
posterior lophs on buccal face. T4 present as well-developed ridge. All M2/ crown
details obliterated by wear. M2/ triangular in outline, with apex facing posteriorly. M/1
heavily worn. Protolophid horseshoe-shaped dentine basin surrounded by remnant of
enamel crown. Hypolophid similar, but subrectangular shaped basin. Posterior cingu-

PROC. LINN. SOC. N.S.W., 111 (3), 1989

T. F FLANNERY Daly)

Fig. I. The crania of A, Microhydromys musseri (holotype) BBM 101737. B, BBM 60202, M. richardsoni.
C, AM M14166, M. richardsonz.

lum large and ovoid, positioned at anterolingual margin of tooth. M/2 smaller than
M/1. Protolophid subrectangular and heavily worn basin. Hypolophid crescent shaped
basin. Lophids separated by simple interlophid valley.

Cranium slightly damaged; left occipital condyle sheared off and basoccipital
shattered then reglued. Rostrum shorter than in other microhydromyines, and
broadened posteriorly. Premaxillae do not extend beyond incisors. Masseteric foramen
lunette shaped and sloping anterodorsally, but with vertical orientation at posterior end.
Infraorbital foramen moderate in size, not expanded as in Hydromys, Parahydromys and
Crossomys species. Incisive foramina short and bowed, posterior ends located just
posteromedial to zygomatic plates. Zygomatic plates broad and flared laterally.
Zygomatic arch thin. Braincase moderately inflated. Bregmatic anteroposteriorly very
short but broad. Palate deeply excavated, with molar alveolar margin raised into a ridge
which extends well ventral to palate floor. Palatal foramina partially obscured by over-
hang of this ridge. Well-defined grooves run from palatal to incisive foramina. Left
pterygoid broken away. Otic notch large, bulla unexpanded.

PROG. LINN. SOG. N.S.W., 111 (3), 1989

218 A NEW MURID FROM PAPUA NEW GUINEA

Fig. 2. Skins of A, M. richardson: (AM M14166) B, ventral view and C, dorsal view of M. richardsoni
(BBM60202), D, ventral view and E, dorsal view of Microhydromys musseri n. sp. (holotype) BBM101737.

DISCUSSION

Systematics

The assignation of the newly described species to the genus Microhydromys is based
upon a cladistic interpretation of aspects of hydromyine morphology. Phenetically,
mussert n. sp. and M. richardsoni appear to be rather different. However they share a
number of key, apparently derived, features that are either rare or are not seen
elsewhere among the Hydromyinae. These include the following.
1) The M1/is extremely narrow and elongate, and more nearly rectangular in occlusal
outline than in other hydromyines. This feature is unique in the subfamily and is not
seen among other Australo-Papuan murids. Thus on the basis of outgroup comparison
this feature 1s likely to be derived.
2) The lingual end of the interloph valley between the anterior and medial lophs of
M1/ is very narrow, and partially closed off by a low ridge from T1. The entire lingual

PROC. LINN. SOC. N.S.W., 111 (3), 1989

T. F FLANNERY 219

margin of the tooth thus forms a continuous linear face which is parallel with the skull
midline. Again, this feature is not seen in other hydromyines and is doubtless derived.

3) The rostrum is short relative to skull length and yet the interorbital canals are not
greatly enlarged. The only other hydromyines with such a short rostrum are the species
of Hydromys, Crossomys and Parahydromys (here called the ‘Hydromys group’). These very
large hydromyines however all possess greatly enlarged infraorbital canals. Most murids
(including the plesiomorphic hydromyine Leptomys elegans) have longer rostra. Thus it
seems likely that the presence of a short skull is a derived feature. Because in other
aspects the morphology of members of the ‘Hydromys group and the species of Micro-
hydromys is so different it seems likely that this similarity is due to convergence.

4) The palatal foramina are partially closed posteriorly and are narrow. Again, these
features are unique in the subfamily and on the basis of outgroup comparison are likely
to be derived.

Other aspects of morphology of the two Maicrohydromys species are however not
shared between these taxa, but are either unique, or are shared by one or the other
species with other hydromyines. These features are interpreted as being either retained
symplesiomorphies or as being due to convergence; the features are as follows. The
pelage of M. mussert n. sp. is bright. Its warm brown and cinnamon tones and sharply
contrasting venter are not approached in any other microhydromyines. Among other
hydromyines, only the species of Leptomys and one species of Paraleptomys possess such
colouration. The species of Leptomys are particularly plesiomorphic hydromyines in
many aspects of their morphology, and no other features would suggest a relationship
with Microhydromys. Thus this feature may well be due to either convergence or else be a
retained symplesiomorphy.

Another striking feature of /. musseri n. sp. that is most closely approached only in
apparently distantly related taxa is the extremely concave palate. Only in Xeromys
myoides, among other hydromyines, is the condition seen in M. musseri n. sp. approached.
However, here details of actual palatal structure differ, suggesting that palatal concavity
in these taxa is the result of convergence. The palatal foramina in X. myozdes are not over-
hung by the ridges supporting the cheekteeth as in M. mussern n. sp., but are subovate
and open. Furthermore, in cross section the palate of M. mussert is more angular than in
X. myoides (where the cross section is arch shaped).

Overall skull shape at first suggested that a close relationship might exist between
M. musserin. sp. and the ‘Hydromys group’ species, for these taxa have a short rostrum and
are superficially similar in skull shape. However, in species of the ‘Hydromys group’ T1
and T4 on M1i/ are not extended posteriorly as they are in other New Guinean
hydromyines. Posterior extension of these cusps must be regarded as derived, as it is not
seen in plesiomorphic murid groups. This suggests that all hydromyines except
members of the ‘Hydromys group may be monophyletic, and that similarities between M.
musseri n. sp. and the species of Hydromys are either plesiomorphic or due to convergence.

A final striking feature of M. musseri n. sp. is the presence of a distinct and large
posterior cingulum on M/1. Only in the species of Leptomys, Paraleptomys and Neohydromys
among hydromyines is this structure otherwise retained. The widespread presence of a
posterior cingulum on the lower molars of other murid groups (including all non-
hydromyine Australo-Papuan taxa) suggests that the loss of this structure is derived.
That the posterior cingulum appears to have been retained in several unrelated
hydromyine taxa indicates that its loss within the subfamily has occurred independently
in several lineages.

Mucrohydromys richardsoni possesses one feature that is unique in the Hydromyinae.
This is the presence of longitudinally grooved incisors. Such grooved incisors are,
however, seen in a number of other murid taxa, such as the species of Mylomys, Pelomys

PROC. LINN. SOC. N.S.W., 111 (3), 1989

220 A NEW MURID FROM PAPUA NEW GUINEA

and Golunda. In Vandeleurta oleracea the incisors are only occasionally grooved, while in
Mus xenodontus only one incisor is grooved. Misonne (1969) suggests that this indicates
that the presence of grooves on the upper incisors of murids is under relatively simple
genetic control and that it has only limited systematic importance. These data, together
with the fact that the grooved incisors of M. mchardsoni are unique (autapomorphic)
among near relatives, suggests that not too much importance should be placed upon this
feature in phylogenetic analysis.

In summary, it would appear that M. musseri n. sp. and M. rchardsoni are each
other’s nearest relatives, but that both possess a number of independent specializations
not seen in the other. This may in turn indicate that these taxa have been evolving
separately for a considerable period of time. On this basis, a case could be made for
either placing each in a monotypic genus, or retaining both in Microhydromys. The latter
course has been chosen because I feel that the recognition of yet another monotypic
genus of microhydromyine murid is not desirable until relationships within the group
are clarified; furthermore such a decision would not indicate the nature of the
relationship between M. richardsoni and M. musseri n. sp.

Biology and Zoogeography

A note on the label of the type specimen of M. musseri n. sp. indicates that it was
taken at an altitude of 1,350m in a snap trap in ‘forest’. At altitudes above about 1,200m
Mt Somoro and the other high peaks of the North Coast Ranges are covered by mossy
forest which differs sharply from the forest at lower altitudes. It is now becoming clear
that these forests harbour a distinctive mammal fauna unlike that found anywhere else
in New Guinea. The first endemic mammal to be described from these ranges (in this
case the Cyclops Mountains) was also a hydromyine, Faraleptomys rufilatus Osgood, 1945.
It has subsequently been found in the Mt Somoro area. Zeigler (1981) described a
second endemic, the large petaurid Petaurus abidi from Mt Somoro. This paper describes
a third endemic, which is yet another hydromyine. Furthermore, recent fieldwork
undertaken in the Somoro area has revealed the presence of additional, as yet undes-
cribed, endemic mammal species. In addition to these taxa, a number of slightly more
widespread species are found in this habitat. These include Pseudocheirops albertisi1, also
known from the Vogelkop and Japen Island, and Dendrolagus inustus, which has a similar
distribution. The area of mossy forest available to these species in the Torricelli Moun-
tains is tiny (ca. 39 km?), but in the recent past was probably larger, when Ice Age
cooling depressed altitudinal zones. However, it appears that at no time in the last few
million years was this region in contact with the mossy forests of the Central Cordillera.
This is reflected in the ‘unbalanced’ assemblage of species in this habitat. For example,
no endemic dasyurids are known from these ranges, while the Central Cordillera has 6
species found exclusively at mid-high altitudes. Perhaps the fact that two out of the three
described endemic mammals from the North Coast Ranges are insectivorous
hydromyines reflects this lack. In the absence of the insectivores and carnivores which
inhabit this forest type elsewhere, it may well have been that the hydromyines that could
invade the habitat underwent a modest radiation. Whatever the case, it is apparent that
no similar event happened elsewhere, as no endemic hydromyine species have been
found on other isolated mountain ranges in New Guinea.

Additional Specimens

The only detailed information previously published regarding Muocrohydromys
richardsoni is that contained in the type description and paraphrases of it. The holotype is
an adult male, taken, presumably in a snap trap (the posterior part of the skull is broken
in a way that suggests snap trap damage), at an altitude of 850m 4km SW of Bernhard

PROC. LINN. SOC. N.S.W,, 111 (3), 1989

T. F. FLANNERY 221

camp on the Idenburg River, Irian Jaya on 16 March 1939. Only three further
specimens have been collected since that time, and two of these have been examined for
this study. The first collected is an adult male (AMNH 198790). This was snap trapped
under a small log in secondary growth at an altitude of 670m at Wanuma in the Adelbert
Mountains, Madang Province by A. C. Zeigler on 21 October 1967. I have been unable
to examine this specimen but Dr G. Musser kindly made the following notes for me; the
terminal 35mm of the tail is mottled white, and the skull is so shattered that measure-
ments could not be made. The Adelbert Range is a small, relatively low mountain range
(most of it barely exceeding 1,000m) which is isolated from the New Guinean Central
Cordillera.

The second additional specimen to be collected 1s another adult male (BBM60202).
This was snap trapped in an area of secondary forest-Eucalypt savannah near Sirinumu
Dam, Sogeri area, Central Province Papua New Guinea on 28 October 1968. Sirinumu
Dam lies at an altitude of ¢. 550m. This is a most unexpected locality, as the Idenburg
River and Adelbert Range specimens must have come from much wetter forest. The
measurements of this specimen closely approach those of the holotype for the most part
(Table 1). The pelage and cranial morphology are also similar to that of the holotype.
They differ primarily in that: approximately 37mm of the distal part of the tail is white
tipped as opposed to 10mm in the holotype; the grey of the dorsum is interrupted by
irregular and inconspicuous ginger blotching (the individual blotches being 1-2mm in
diameter).

TABLE 1
Measurements for the 5 known specimens of Microhydromys. Measurements for AMNH 152079 are from Tate (1951).
Measurements in mm and weight in grams. 1 = length, w = width, inc. = incisive
M. mussert M. richardsoni
Measurement BBM-NG AMNH AMNH AMM BBM-NG
/Weight 101737 198790 152079 14166 60202
head body 1. 108 83 80 86 86
tail vent 1. 101 84 92 79 83
hindfoot (su) 1. 22 19 20 = 19
ear (n) 1. 13 11 8 = 12
weight — 9 = 12 =
condylobasal 1. 23.6 = = 20.2 19.6
bizygomatic w. i133}.40 = Q2 10.1 9.6
palate 1. 13.6 = 10.0 9.5 Q7
rostral w. 5.7 — = 4.1 4.0
inc. foramen 1. DP = ES) 2.1 1.9
interorbital w. D2 — 4.4 4.6 4.2
nasal 1. 8.0 = 6.4 7.0 5.9
maximum nasal
W. 3.0 - 2.6 2.6 223
mastoid w. 10.7 = 9.4 8.8 Or
bulla 1. 4.4 = 3.8 Aa? At)
Mi1-2/ 1. 29) = 2.4 2.0 1.9
M1-1/ w. internal 3.5 = 2.4 2.7 Ped)
M1/ 1. Dell — 1.6 1.4 1.2
M1/ w. 0.9 = 0.8 0.8 0.6
M2/ 1. 0.8 = 0.8 0.7 0.7

M2/ w. 0.8 = 0.7 0.6 0.6

The most recently collected specimen is AM M14166. It is an adult female snap
trapped on 18 September 1979 at 1,450m on the southern slopes of Mt Sisa, Southern
Highlands Province by P. Dwyer. Although most measurements are similar to those for
the other M. richardsoni individuals known, it differs in a number of ways. The nasals are

PROC. LINN. SOG. N.S.W., 111 (3), 1989

DY A NEW MURID FROM PAPUA NEW GUINEA

longer and the mastoid width narrower than in the male specimens (Table 1). The palate
does not extend as far posteriorly behind M2/ as in the Sogeri specimen. The skin of the
Mt Sisa specimen is flat. It closely resembles that of the Sogeri individual except that it
lacks the ginger blotching and has a small patch of pure white hairs on the throat. The
most distal 25mm of the tail is pure white, while a light and dark mottled area extends a
further 30mm proximally.

It is unclear as yet whether the differences between AM M14166 and the other M.
richardsoni specimens represent sexual dimorphism, intra, or possibly even interspecific
variation. However from the specimens described here it is clear that this species, or taxa
closely related to it, are relatively widely distributed across New Guinea, both
geographically and in differing environments.

CONCLUSION

Microhydromys musseri n. sp. is described. Known from a single specimen collected in
mossy forest high in the Torricelli Mountains, it differs in many aspects of its mor-
phology from M. richardson. Three additional specimens of M. richardsoni are described.
They indicate that this species has a wide geographic distribution and that it exhibits
considerable morphological variation.’ Microhydromys richardsont can inhabit drier
habitats than had previously been suspected.

ACKNOWLEDGEMENTS

In particular I would like to thank Dr Guy Musser of the American Museum of
Natural History for his invaluable comments upon an earlier draft of this paper, and for
supplying details of the Microhydromys specimens held in the American Museum. Mr
Ken Aplin of the University of New South Wales also kindly commented upon this
manuscript and made many useful suggestions. I would also like to thank Dr Allan
Allison of the Bishop Museum for making the hydromyine specimens held in the Bishop
Museum, Hawaii, available for me to study, also Mr Alan Ziegler for his help in this
matter. Ms Tish Ennis produced the figures.

References
MENZIES, J. and DENNIS, E., 1979. — Handbook of New Guinea Rodents. Wau Ecology Institute Handbook No.
6.
MISONNE, X., 1969. — African and Indo-Australian muridae. Evolutionary trends. Annis Mus. r. Afr. cent.
172: 1-219.
MUSSER, G., 1981. — The giant rat of Flores and its relatives east of Borneo and Bali. Bull. Amer. Mus. Hist.
169: 69-175.

MUSSER, G., 1982. — Results of the Archbold Expeditions. No. 110. Crunomys and the small-bodied shrew
rats native to the Philippine Islands and Sulawesi (Celebes). Bull. Amer. Mus. Nat. Hist. 174: 1-95.

OsGooD, W. H., 1945. — A new rodent from Dutch New Guinea. Fieldiana (Zoology) 31: 1-2.

SMITHE, F. B., 1974. — Naturalist’s Colour Guide Supplement. New York: American Museum of Natural History.

Tate, G. H. H., 1951. — Results of the Archbold Expeditions. No. 65. The rodents of Australia and New
Guinea. Bull. Amer. Mus. Nat. Hist. 97: 183-430.

Tare, G. H. H. and ARCHBOLD, R., 1941. — Results of the Archbold Expeditions. No. 31. New rodents and
marsupials from New Guinea. Amer. Mus. Novit. 1101.

THOMAS, O., 1898. — On the mammals obtained by Mr John Whitehead during his recent expedition to the
Philippines. Trans. Zool. Soc. London 14: 377-412.

ZIEGLER A. C., 1981. — Petaurus abidi, a new species of glider (Marsupialia, Petauridae) from Papua New
Guinea. Aust. Mammal. 4: 81-88.

PROG. LINN. SOG. N.S.W., 111 (3), 1989

PROCEEDINGS
of the

LINNEAN
SOCIETY

NEW SOUTH WALES

VOLUME 111
NUMBER 4

Tidal and Diel Variations in the Abundance of
Larval Fishes in Botany Bay, New South Wales,
with Emphasis on Larval Silverbiddy Gerres ovatus

(Fam. Gerreidae) and Gobies (Fam. Gobiidae)

ALDO S. STEFFE

STEFFE, A. S. Tidal and diel variations in the abundance of larval fishes in Botany Bay,

New South Walcs, with emphasis on larval silverbiddy Gerres ovatus (Fam.
Gerreidac) and gobies (Fam. Gobiidae). Proc. Linn. Soc. N.S.W. 111 (4), 1989:
225-232.

Surface plankton samples were collected from an area of strong tidal flow in
Botany Bay, during early autumn 1981, to examine tidal and diel variations in the
abundance of larval fishes. The sampling program was restricted to one 24 hour period,
and yielded 2,898 larvae consisting of 30 distinct larval types from 21 families. Larval
gobies (Fam. Gobiidae) and silverbiddy Gerres ovatus (Fam. Gerreidae) dominated the
assemblage, and together accounted for 82.5% of the standardized total catch. On the
selected sampling day the composition of the larval assemblage differed at cither end of
the tidal range. Gobiid larvae were more abundant at low tide both day and night,
whereas abundances of G. ovatus were greater at high tide both day and night. Night
catches of gobiid and G. ovatus larvae were greater than day catches. The great majority
of G. ovatus larvae had deflated gas bladders during the day and inflated gas bladders at
night. The limitations of these findings are acknowledged; due to the short sampling
period it is possible that effects of other variables were confounded with effects of the
main factors being tested.

Aldo S. Steffe, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109;
manuscript recewed 15 March 1989, accepted for publication 19 July 1989.

INTRODUCTION

Although the nursery function of Australian estuaries has been studied (Lenanton,
1977; Robertson, 1980; SPCC, 1981; Young, 1981; Bell et al., 1984; Middleton et al., 1984)
little is known about larval fish distributions or the factors which affect larval recruit-
ment in these estuarine areas (Steffe and Pease, 1988).

The common problem facing both recruiting larvae and those already in the
estuary is how to maintain their position in the estuary and avoid being flushed out on
the ebb tide. Weinstein et al. (1980) proposed that successful recruitment and/or
retention in a stratified estuarine system involved selective vertical migrations by larval
fishes in conjunction with changes in tide and photoperiod. There is also evidence that
the larvae of other estuarine dependent species respond to tidal or diel stimuli, or both
(Fore and Baxter, 1972; Graham, 1972; Eldridge, 1977; Melville-Smith et al., 1981;
Fortier and Leggett, 1982; Norcross and Shaw, 1984; Roper, 1986), but the mechan-
ism(s) by which larval fishes achieve this are poorly understood.

Here, the findings of a sampling program designed to analyse variation in larval
fish abundances with respect to changes in tidal and diel conditions are reported. The
main question asked was: on the selected sampling day did larval fish abundances at the
surface vary with respect to changes in tide and/or diel conditions?

MATERIALS AND METHODS

Study Area
Botany Bay (34°01’S, 151°11’E) is a large, semi-landlocked estuary on the east
coast of Australia (Fig. 1). It is dominated by ocean swell and wind waves (Roy e¢ al.,

PROG. LINN. SOC. N.S.W., 111 (4), 1989

226 LARVAL FISHES IN BOTANY BAY

1980), is vertically well mixed (Rochford, 1951), and at most times is best described as a
marine embayment.

Fig. 1. Map of Botany Bay showing location of sampling station. Abbrev.: TP = Towra Point.

A sampling station (Fig. 1) was selected off Towra Point because of its position in
the main tidal stream and its close proximity to large Poszdonia australis and Zostera
capricorn’ seagrass meadows, and an extensive mangrove stand. These habitats are
important fish nursery areas (Bell e¢ al., 1984; Middleton et al., 1984). Samples were
collected during March (early Autumn) as juveniles of many economically valuable
fishes are most abundant in the Bay shortly after this period (Bell, 1980; SPCC, 1981;
Bell et al., 1984; Middleton et al., 1984). The maximum depth at the sampling station was
about 4m.

Field Procedures

The sampling program was designed to analyse variances in larval fish numbers
between stage of tide (high vs low) and diel condition (day vs night) at one station. The
design was orthogonal and required that both high and low tides be sampled during day-
light, and at night. High and low tide were defined as a two hour period with the tidal
prediction at its centre. Four replicate samples were collected at each consecutive high
and low tide over the 24 hour sampling period. Tidal height ranged from 0.2-1.6 m above
I.S.L.W. during the sampling period.

Day sampling commenced 40 minutes after sunrise (05.57h) on 19 March 1981 and

PROC. LINN. SOC. N.S.W., 111 (4), 1989

A. S. STEFFE Dah

night sampling commenced one hour after sunset (18.09h). Plankton samples were
collected from a small (4m) boat using a net with a square mouth (area 0.25m?’), a mesh
of 500 wm, and a length of about 2.5m. The net was towed in a circular path at the
surface at a constant speed of about 2m sec? for 5 minutes (+ 5 sec.). This procedure
kept the net out of the engine wash and thus avoided unnecessarily increasing net
escapement. A General Oceanics (model 2030) flowmeter was used to measure the
volume of water filtered. Volume filtered per tow averaged 133.7m/? (s.d. 13.2). A salinity
(+ 0.1°/0c) and water temperature (+ 0.2°C) reading was taken during each of the
consecutive tides sampled.

Samples were preserved immediately after collection in 5% Steedman’s preser-
vative and were sorted entirely under a dissecting microscope. All larval fishes were
identified to the lowest possible taxonomic level and then stored in 4% buffered forma-
lin. Standardized larval abundances are expressed as the number of larvae per 100m?
water filtered. Terminology follows the definitions of Leis and Rennis (1983).

Size Distribution and Gas Bladder Inflation Incidence

Some 200 day-caught and 200 night-caught larvae of Gerres ovatus and of gobiids
were randomly selected. These larvae were measured to the nearest 0.1 mm using a
dissecting microscope with mounted ocular micrometer. Notochord length was
recorded for preflexion and flexion stages whilst standard length was measured for post-
flexion stages. The numbers of larvae with deflated, or inflated gas bladders were then
recorded separately for day-caught and night-caught G. ovatus only. This was easily done
as the gas bladder in this species is clearly visible when inflated. This procedure was not
repeated for gobiid larvae as they have a permanently inflated gas bladder.

RESULTS

The sampling program yielded 2,898 larval fishes consisting of 30 distinct larval
types from 21 families (Table 1). Silverbiddy Gerres ovatus and gobiid larvae dominated
the larval fish assemblage. G. ovatus larvae accounted for 31.3% of the standardized total
catch whilst gobiids (four spp.) made up 51.2% of this total. The remaining 19 families
contributed 17.5% of the standardized total catch but were not sufficiently abundant, or
were present in too few samples, to warrant statistical analysis (Table 1).

More Gerres ovatus were caught at night and during high tide (Fig. 2a, Table 2). The
interaction term was not significant. Similarly, significantly more gobiid larvae were
caught at night, however unlike Gerres ovatus, significantly more gobiids were caught
during low tide (Fig. 2b, Table 2). There was no significant interaction (Table 2).

Total fish larvae reflected the contrasting patterns of Gerres ovatus and gobiids. Night
catches were significantly greater than day catches, but there was no significant tidal
effect or interaction (Fig. 2c, Table 2).

The length frequency of night-caught Gerres ovatus larvae was similar to that for day
caught larvae (Kolmogorov-Smirnov test, Dmax. = 0.075, p> >>0.05) (Fig. 3a).
Ninety-eight percent of day-caught larvae (n = 200) were found to have deflated gas
bladders and ninety-five percent of night-caught larvae (n = 200) had strongly inflated
gas bladders.

The length frequency distribution of gobiid larvae caught during daylight was
significantly different to that of gobiids caught at night (Kolmogorov-Smirnov test,
Dmax. = 0.23, p<0.001) (Fig. 3b). In contrast to G. ovatus, more larger gobiid larvae
were taken at night.

Salinity and water temperature did not fluctuate greatly during sampling (temp.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

228 LARVAL FISHES IN BOTANY BAY

22.8-24.0°C; sal. 32.7-34.0°/o0) and rainfall had not been reported for the previous ten
days in the area (Bureau of Meteorology, 1981).

TABLE 1

Number of distinct larval types, occurrence, and the percentage of the standardized total catch for each taxon in the larval
assemblage. Note that each larval type may not be monospecific

No. Larval Occurrence % Standardized

‘Taxon ‘Types (Max. = 16) total catch
Gobiidac 4 16 51.2
Gerreidac | 16 31.3
Ambassidae 2 15 4.7
Syngnathidac 4 16 3.7
Blenniidae 2 12 1.5
Sillaginidae 1 8 0.7
Sparidac 2 7 0.7
Monacanthidac 1 7 0.5
Carangidac | 5 0.4
Atherinidac 1 4 0.3
Anguilliformes 1 3 0.2
Clupeidac 1 5 0.2
Hemirhamphidac 1 1 0.1
Platycephalidae 1 1 0.1
Pempherididac 1 3 0.1
Soleidac 1 3 0.1
Tetraodontidac 1 3 0.1
Scorpaenidac 1 1

Mugilidac 1 1

Sphyraenidac 1 1 0.1
Callionymidac 1 1

Damaged larvac = 16 4.0

TABLE 2

F ratios and significance levels derived from two way fixed effects ANOVA of effects of tade (high vs low) and diel period (day vs
night) for Gerres ovatus, Gobiidae and sample totals. Data tested for heteroscedasticity using Cochran’ test (p< 0.1). A =
log transformed data, B = raw data

F ratio
Source of Gerres Sample
Variance df ovatus’ Gobiidac® Totals?
Tide 1 16.10** 5.7 0.08 NS
Dicl 1 1D. So" 19) WSS" il le
TxD 1 1.85 NS 0.06 NS 0.08 NS
Residual 12

*» <0.05, ** p <0.01, *** p <0.001, NS p >0.05

DISCUSSION

Gerres ovatus larvae were significantly more abundant on high tides whilst gobuid
larvae were caught in significantly greater numbers during low tides. As G. ovatus and
numerous gobiid species are known to spawn within Botany Bay (State Pollution
Control Commission, 1981) their centres of larval abundance may have been expected to
coincide. Yet, they have become spatially separated, occurring at either end of the tidal
range. This larval distribution was found both during daylight and at night. Two
hypotheses, not mutually exclusive, to explain this are: (1) spawning aggregations of G.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

(a)
100

S) 80

a 60

= 40

—

= 20

NIGHT

HIGH

TIDAL STAGE

(c)

N@ LARVAE PER 100m3

250

200

150

100

AGS) SBE EE 229

160
140
wae NIGHT
100

NO LARVAE PER 100m?

HIGH LOW
TIDAL STAGE

NIGHT

DAY

HIGH LOW
TIDAL STAGE

Fig. 2. Mean and + one SE for the standardized larval abundances at each of the consecutive tides, both day
and night, over the 24 hour sampling period for (a) Gerves ovatus, (b) Gobiidae, and (c) sample totals.

PROG. LINN. SOC. N.S.W., 111 (4), 1989

230 LARVAL FISHES IN BOTANY BAY

(a)
(b)

pce eew em eww ew een’

FREQUENCY
FREQUENCY

Bosooosoo00K

DTS Ran eS IGS 7 123. > 42 35006
LENGTH (mm) LENGTH (mm)

Fig. 3. Length frequencies of randomly selected day-caught (n = 200) and night-caught (n = 200) larvae for
(a) Gerres ovatus, and (b) Gobiidae. Solid lines denote day-caught, and broken lines denote night-caught.

ovatus occur near the Bay entrance and seaward of gobiid spawning sites; and (2) both
groups spawn at a similar distance into the Bay, but the demersal eggs of gobiids are
more effective than the pelagic eggs of G. ovatus at reducing the net transport of offspring
seaward from spawning sites by currents. This could occur because demersal eggs,
unlike pelagic eggs, are not subject to passive transportation by currents, and because at
hatching, larvae from demersal eggs tend to be relatively larger, more developed, with
better swimming capabilities than those from pelagic eggs (Steffe and Pease, 1988).
Larval catches of Gerres ovatus and gobiids were found to be significantly greater at
night (Table 2). There was no difference in the size structure of G. ovatus between day
and night (Fig. 3a) suggesting that differential net avoidance was minimal and that the
higher night catches could be mainly attributed to the effects of diel vertical migration.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

AS. SUEFFE 231

The effects of differential net avoidance and diel vertical migration could not be
separated for gobiids as more large larvae were caught at night (Fig. 3b).

Most Gerres ovatus larvae had deflated gas bladders during the day whilst the reverse
was true at night. This phenomenon appears to be common and occurs in many taxo-
nomically diverse teleost groups (Hunter and Sanchez, 1976; Leis and Rennis, 1983;
Liew, 1983; Hoss and Phonlor, 1984; Kitajima et al., 1985; A. Steffe, unpub. data).
Hunter and Sanchez (1976) found that nocturnal gas bladder inflation can provide
considerable energy savings to northern anchovy Engraulis mordax larvae by retarding
sinking. It is likely that larval G. ovatus gain a similar benefit.

The silverbiddy, G. ovatus, has been found to recruit almost exclusively to mangrove
areas in Botany Bay (State Pollution Control Commission, 1981). The observed larval
distribution of G. ovatus at a site near its preferred nursery habitat supports the hypothe-
sis that larval silverbiddy may be using flood tides to assist their transportation into
mangrove areas. This hypothesis appears tenable because mangrove areas are wholly
dependent on tide for water exchange.

The data presented and interpreted here are based on collections made during a
single 24 hour period. Consequently, it is possible that the effects of other variables
which were not specifically tested may have, by chance, been confounded with the effects
of the tidal and diel factors examined during the selected sampling day; in view of the
relatively constant temperature and salinity conditions it is unlikely that larval catches
were influenced by these parameters. Further sampling, that is, the same experiment
repeated on other ‘replicate’ days, is required to eliminate this possible source of error
and to possibly allow the conclusions drawn here to be accepted with greater confidence.

ACKNOWLEDGEMENTS

I thank B. Griffiths, B. Hodgson, J. Leis, J. MacIntyre, A. Mazanov, A.
Miskiewicz and M. Westoby all of whom contributed to the compilation and completion
of this work. Special thanks to my parents and wife for financial support and field
assistance. L. Beckley, J. Bell, W. Gladstone, J. Leis, K. McGuiness, and M. Westoby
made useful suggestions which improved the manuscript.

References

BELL, J. D., 1980. — Aspects of the ecology of fourteen economically important fish species in Botany Bay,
New South Wales, with special emphasis on habitat utilization and discussion of the effects of man-
induced habitat changes. North Ryde, N.S.W.: Macquarie University, M.Sc. thesis, unpubl.

— , POLLARD, D. A., BURCHMORE, J. J., PEASE, B. C., and MIDDLETON, M. J., 1984. — Structure ofa fish
community in a temperate tidal mangrove creek in Botany Bay, New South Wales. Aust. J. Mar. Freshw.

Res. 35: 33-46.

BUREAU OF METEOROLOGY, 1981. — Monthly weather review. New South Wales. Dept. of Science,
Canberra.

ELDRIDGE, M. B., 1977. — Factors influencing distribution of fish eggs and larvae over eight 24 hour

samplings in Richardson Bay, California. Calif. Fish and Game 63: 101-116.

Fore, P. L., and BAXTER, K. N., 1972. — Diel fluctuations in the catch of larval gulf menhaden, Brevoortia
patronus, at Galveston Entrance, ‘lexas. Trans. Amer. Fish. Soc. 101: 729-732.

Fortier, L., and LEGGETT, W. C., 1982. — Fickian transport and the dispersal of fish larvae in estuaries.
Canadian J. Fish. Aqu. Sci. 39: 1150-1163.

GRAHAM, J. J., 1972. — Retention of larval herring within the Sheepscot estuary of Maine. U.S. Fish. Bull.

70: 299-305.
Hoss, D. E., and PHONLOR, G., 1984. — Field and laboratory observations on diurnal swim bladder
inflation-deflation in larvae of gulf menhaden, Brevoortza patronus. U.S. Fish. Bull. 82: 513-517.
HUNTER, J. R., and SANCHEZ, C., 1976. — Diel changes in swim-bladder inflation of the larvae of the

northern anchovy, Engraulis mordax Girard. U.S. Fish. Bull. 74: 847-855.
KiTajIMA, C., TSUKASHIMA, Y., and TANAKA, M., 1985 — The voluminal changes of swim bladder of larval
red sea bream Pagrus major. Bull. Japan. Soc. Sci. Fish. 51: 759-764.

PROC. LINN. SOG. N.S.W., 111 (4), 1989

DBD LARVAL FISHES IN BOTANY BAY

Leis, J. M., and RENNIS, D. S., 1983. — The larvae of Indo-Pacific coral reef fishes. Kensington, N.S.W.: Univer-
sity N.S.W. Press and Honolulu: University of Hawaii Press.

LENANTON, R. C. J., 1977. — Aspects of the ecology of fish and commercial crustaceans of the Blackwood
River estuary, Western Australia. Fish. Res. Bull. WA. 19: 1-72.

Liew, H. C., 1983. — Studies on flatfish larvae (Fam. Psettodidae and Bothidae, Pleuronectiformes) in the
shelf waters of the central Great Barrier Reef, Australia. Townsville, Qld.: James Cook University,
M.Sc. thesis, unpubl.

MELVILLE-SMITH, R., BAIRD, D., and WOOLDRIDGE, T., 1981. — The utilization of tidal currents by the
larvae of an estuarine fish. Sth. African J. Zool. 16: 10-13.

MIDDLETON, M. J., BELL, J. D., BURCHMORE, J. J., POLLARD, D. A., and PEASE, B. C., 1984. — Structural
differences in the fish communities of Zostera capricornt and Posidonia australis seagrass meadows in
Botany Bay, New South Wales. Aquat. Bot. 18: 89-109.

Norcross, B. L., and SHAW, R. F., 1984. — Oceanic and estuarine transport of fish eggs and larvae: a
review. Trans. Amer. Fish. Soc. 113: 153-165.

ROBERTSON, A. I., 1980. — The structure and organization of an eelgrass fish fauna. Oecologia 47: 76-82.

ROCHFORD, D. J., 1951. — Studies in Australian estuarine hydrology. I. Introductory and comparative
features. Aust. J. Mar. Freshw. Res. 2: 1-116.

Roper, D. S., 1986. — Occurrence and recruitment of fish larvae in a northern New Zealand estuary. Est.
Coast. Shelf Scr. 22: 705-717.

Roy, P. S., THoM, B. G., and WRIGHT, L. D., 1980. — Holocene sequences on an embayed high-energy
coast: an evolutionary model. Sedimentary Geology 26: 1-19.

STATE POLLUTION CONTROL COMMISSION, 1981. — The ecology of fish in Botany Bay — biology of commer-
cially and recreationally valuable species. SPCC Rep. No. BBS 23B, 1-287.

STEFFE, A. S., and PEASE, B. C., 1988. — Diurnal survey of ichthyoplankton abundance, distribution and
seasonality in Botany Bay, New South Wales. Proc. Linn. Soc. N.S.W. 110: 1-10.

WEINSTEIN, M. P., WEISS, S. L., HODSON, R. G., and GERRY, L. R., 1980. — Retention of three taxa of post-
larval fishes in an intensively flushed tidal estuary, Cape Fear River, North Carolina. U.S. Fish. Bull.
78: 419-436.

YOUNG, P. C., 1981. — Temporal changes in the vagile epibenthic fauna of two seagrass meadows (Zostera
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PROC. LINN. SOC. N.S.W., 111 (4), 1989

Redescription of Austrochthonius australis Hoff
(Chthonidae: Pseudoscorpionida)

CLARICE M. A. KENNEDY

KENNEDY, C. M. A. Redescription of Austrochthonius australis Hoff (Chthoniidae:
Pseudoscorpionida). Proc. Linn. Soc. N.S.W. 111 (4), 1989: 233-240.

An examination of approximately 150 specimens of Austrochthonius australis Hoff
from various locations in southeastern Australia suggests a considerable size variation
within the species when compared with the two published descriptions of adults (Hoff,
1951; Beier, 1966). For the first time the nymphal stages are described.

Clarice M. A. Kennedy, School of Biological Sciences, Macquarie University, North Ryde, Aus-
tralia, 2109, manuscript received 7 February 1989, accepted for publication 19 July 1989.

INTRODUCTION

A. australis 1s widely distributed in a variety of habitats throughout Australia. The
description of the holotype is based on a single female specimen from Mt Slide, |
Victoria. Beier (1966) has briefly described male and female specimens based on an
examination of material from Western Australia, Victoria, New South Wales,
Australian Capital Territory and Tasmania.

Investigation of specimens from Western Australia, identified as A. australis, has
revealed a number of differences from the holotype suggesting that an undescribed
species is represented. In addition, individuals of a population in the Sydney region are
much smaller, when compared with the two published descriptions of the adult by Hoff
(1951) and Beier (1966); this prompted examination of some 150 specimens from various
locations in Victoria, New South Wales and Tasmania to determine if specimens identi-
fied as A. australis are in fact a single species. Apart from size, specimens examined are
morphologically similar to the holotype.

The results of this research are presented below and represent a more compre-
hensive description of the species which for the first time, incorporates the nymphal
stages.

MATERIALS AND METHODS

Specimens were borrowed from the following institutions: American Museum of
Natural History, New York; Australian Museum, Sydney; Museum of Victoria,
Melbourne and 34 samples from the private collection of Dr. M. S. Harvey, Western
Australian Museum, Perth.

Measurements are based on the examination of at least 5-15 adults of each sex from
various locations in south eastern Australia made in accordance with those described by
Chamberlin (1931). Figures in parentheses are female values and follow those of the
male. The morphological nymphal information is based on measurements of 10
specimens of each stage collected from the Sydney region. Nymphs from the other
regions within the distribution of the species are, at present, unavailable for
comparison.

Abbreviations for chelal trichobothria and setal formulae follow those devised by
Chamberlin (1931). Genitalia terminology follows that of Legg (1975). Specimens of A.
australis from the Sydney region are deposited in the Australian Museum, 6 © (KS
20186), 6 9 (KS 20187), 6 nymphs (KS 20188).

PROG. LINN. SOC. N.S.W,, 111 (4), 1989

234 REDESCRIPTION OF PSEUDOSCOR PION

SYSTEMATIC DESCRIPTION
CHTHONIIDAE Hansen 1893
Austrochthonius Chamberlin 1929

Type species: Chthonius chilensis Chamberlin 1923

Austrochthonius australis Hotf

(Figs 1-4)

Diagnosis: Chelicera with 5 setae on hand; palpal teeth quadrate and separated; 6 pin-
nate coxal spines present on coxa 2; intercoxal tubercle absent. L/W ratio of chela (with
pedicel) range 1:1.05-1.33 (1: 1.03-1.45) x longer than broad.

Description: Adults. Colour light golden brown; carapace surface smooth; pleural
membrane with series of irregularly spaced longitudinal ridges, each with minute,
elevated, white nodules (Fig. 1A). Pedipalp: trochanter stout, femur with lateral
margins virtually parallel, abruptly converging distally, L/W ratio range 1: 1.00-1.65 (1:

VOam . ; g . | VOmpm

Fig. 1. Austrochthonius australis Hoff, scanning electron micrographs, male: A, pleural membrane; B,
chelicerac, dorsal aspect; C, right cheliccra movable finger, ventral aspect; D, left chelicera, serrula exterior
proximal blades; E, right chelicera, scrrula exterior distal blades, ventral aspect; F, right chelicera, scrrula
exterior distal blades, dorsal aspect.

PROC. LINN. SOG. N.S.W., 111 (4), 1989

C. M. A. KENNEDY D335)

1.00-1.65), tibia calyciform; chelal hand broad and shorter than chelal fingers, lateral
margins very slightly rounded, poorly developed pedicel, L/W ratio of chela (with
pedicel) range 1: 1.05-1.33 (1: 1.03-1.45), chela (without pedicel) 1: 1.03-1.31 (1: 1.01-1.42)
x longer than broad. Chelal fingers long and tapering, fixed finger with 8 trichobothria,
26 and zsh medial on dorsum of chelal hand; movable chelal finger with 4 trichobothria;
marginal teeth quadrate, separated and retrorse on fixed finger, teeth smaller on
movable finger becoming broader with rounded apices and laterally fused proximally,
46-50 (0) 45-46 (9 ) (Fig. 4T); venom apparatus absent. Chelicera large, stout with 5
setae on hand, fingers cross distally (Figs 1B, 4S), fixed finger with reticulate sculpture,
teeth 15-16 (0”) 15-17 (Q ), distal one large, remainder diminishing in size proximally;
movable finger with 18 well-spaced smaller teeth which become almost indistinguish-
able proximally; lamina exterior absent. Serrula exterior detached in upper half with 14-

Fig. 2. Austrochthonius australis Hoff, scanning electron micrographs, male: G, right chelicera, flagellum blades,
ventral aspect; H, female, right chelicera showing galea a sclerotic tubercle; I, carapace; J, male, epistome;
K, coxal spines; L, enlargement of pinnate coxal spine blades.

15 (0°) 14(Q ) lamellae (Fig. 1C), lamellae relatively broad, elongate, with parallel sides
proximally, terminally each blade digitate (Fig. 1D) becoming broad distally (Figs 1E,

PROC. LINN. SOC. N.S.W., 111 (4), 1989

236 REDESCRIPTION OF PSEUDOSCORPION

Fig. 3. Austrochthonius australis Hoff, scanning electron micrographs: M, male genital region, external aspect;
N, female genital region, external aspect; P, male genitalia, internal aspect; Q, female genitalia, internal
aspect.

F); galea a sclerotic tubercle in 9 (Fig. 2H), absent in ©’; galea seta basad of mid-point
of movable finger. Flagellum with 8 stalk-like blades each arising separately along a
diagonal slit basad of fixed cheliceral finger (Fig. 1C) becoming unipinnate distally (Fig.
2G). Carapace: subquadrate, surface smooth; anterior margin slightly serrate laterally
with prominent, blunt, dentate epistome (©’) (Fig. 2J), more or less acute (@ ), weakly
reticulate sculpture; lateral margins distinctly narrowed posteriorly (Fig. 21); posterior
margin relatively straight; setae long, acuminate 4: 2: 12 (0), 4: 2: 12-14 (2 ); L/W ratio

PROG. LINN. SOG. N.S.W., 111 (4), 1989

Cc. M. A. KENNEDY Zh

(based on ocular width) range 1: 1.00-1.13 (1: 1.03-1.36), (based on posterior width) 1:
1.07-1.52 (1: 1.12-1.33) x longer than broad. Anterior eye well developed, posterior eye
flattened disc. Tergal chaetotaxy: setae relatively long, acuminate, O 4: 4: 3-4: 2-4: 6:
Goiioa a Oa) Oa Ae ee O07 014-0: 2. UinIsenlate. Goxalichaetotaxya 6
pinnate coxal spines placed anterior-ventrally along proximal half of coxa 2 (Figs 2K, L)
Oe eae srl O a DO 23-45 One 4) 2:92 1-2)02.3 2. Ons nO: 2-32.45. 102 2213)
Intercoxal tubercle absent. Male genitalia (Fig. 3P): ejaculatory canal atrium (ejca)
pear-shaped; roof of each medial diverticulum thickened (trmd) to form two elongated
structures, anterior horns of each free but fused posteriorly, in mid-line is atrium of
posterior dorsal gland (apdg). Cuticle of genital atrium thickened and extended to form
apodemes; dorsal apodeme (da) well developed with Y-shaped structure below ejacu-
latory canal atrium (ejca), posterior extension of Y forms anterior crest of dorsal
apodeme (acda), posterior to atrium of posterior dorsal gland (apdg) the posterior crest
of the dorsal apodeme (pcda) arises in mid-line above duct of median genital sac; lateral
apodemes (la) reduced and thickened to provide attachment for muscles of ejaculatory
canal and support for anterior region of genital region of genital atrium. Chaetotaxy:
anterior operculum with 8-9 relatively long setae along anterior margin, 4-5 along
posterior margin which is deeply concave medially; posterior operculum with 8
relatively long acuminate setae on each side of a deep median notch. These setae project
into the centre forming a grill-like pattern across the genital atrium (Fig. 3M). Female
genitalia (Fig. 3Q): lateral cribriform plates (Icp) clearly differentiated with numerous
pores; median cribriform plate (mcp) — pores scattered over the surface with some
aggregations. Chaetotaxy: anterior operculum a triangular plate; posterior operculum a
simple elongate plate (Fig. 3N). Sternites with reticulate sculpture, chaetotaxy: O 0: 13-
dee Wa /2 dee Gee Saas Ge (Oe (OR 4-153 Ges 23 © (Os ile We os (Os 403 (3 Os Os Oe A, Seizie Nora.
acuminate. Sternites 4-11 uniseriate.

Dimensions (mm): body length 0.75-117 (0.77-1.62); pedipalp: femur 0.21-
0.36/0.07-0.10 (0.21-0.46/0.07-0.10), chela (with pedicel) 0.43-0.61/0.09-0.13 (0.45-
0.70/0.10-0.16), chela (without pedicel) 0.41-0.59/0.09-0.13 (0.41-0.68/0.10-0.16),
movable finger length 0.25-0.37 (0.28-0.45). Carapace 0.27-0.37/ocular width 0.27-
0.41/posterior width 0.21-0.34 (0.28-0.46/ocular width 0.30-0.48/posterior width
0.26-0.43).

Tritonymph. L/W ratio: pedipalpal trochanter 1.00-2.00, femur 2.28-3.00, tibia
1.28-1.42, chela (with pedicel) 3.55-4.33, chela (without pedicel) 3.33-4.11 x longer than
broad. Fixed finger with 7 trichobothria, movable finger with 3 trichobothria, zsb and sb
absent; serrula exterior of chelicera with 11-12 broad, elongate lamellae, each blade
digitate terminally. Carapace: anterior margin slightly serrate laterally, epistomal
process prominent; posterior margin relatively straight; lateral margin narrowing at
point of union with abdominal segments, 4: 2: 8, setae long, acuminate. L/W ratio
(based on ocular width) 1.08-1.25, (based on posterior width) 0.96-1.00 x longer than
broad. Tergal chaetotaxy: 4: 4: 4: 4: 4-6: 4-6: 6: 6: 6: 4: 4-6: 2. Sternal chaetotaxy: 0: 3-4:
6-7: 6: 6: 4: 4: 6: 6: 6-7: 5-6: 2. Coxal chaetotaxy: 4-5 coxal spines in medial position
Alone antcHonimaeimon2nd coxa, 22) 11023010: 302-35 Onl 2-45 Olena

Dimensions (mm): body length 0.66-0.79; pedipalp: trochanter 0.07-0.09/0.05-
0.07, femur 0.16-0.21/0.07, tibia 0.09-0.10/0.07, chela (with pedicel) 0.32-0.39/0.07-0.09,
chela (without pedicel) 0.30-0.37/0.07-0.09, movable finger length 0.19-0.27; carapace
0.27-0.30/ocular width 0.27-0.31/posterior width 0.22-0.25.

Deutonymph. L/W ratio: pedipalpal trochanter 1.20-1.40, femur 2.00-3.20, tibia
1.00-1.75, chela (with pedicel) 3.75-4.50, chela (without pedicel) 3.28-4.16 x longer than
broad. Fixed finger with 6 trichobothria, movable finger with 2 trichobothria, zsd, esd, sb,
6 absent; serrula exterior of chelicera with 9-10 broad, elongate lamellae, each blade

PROC. LINN. SOC. N.S.W., 111 (4), 1989

238 REDESCRIPTION OF PSEUDOSCORPION

ee!

0.14¢m

Fig. 4. Austrochthonius australis Hoff, male: S, right chelicera, dorsal aspect; T, right chela, lateral aspect; Ta,
inset, enlargement showing dentition type (not to scale); U, first right leg; V, fourth right leg; W, left pedipalp,
dorsal aspect.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

Cc. M. A. KENNEDY 239

digitate terminally. Carapace: anterior margin serrate, epistome prominent; 4: 2: 8: 3-6
relatively long setae, 1.05-1.18 x longer than broad (ocular and posterior width equal).
Tergal chaetotaxy: 4: 4: 4: 2-4: 4-6: 4-6: 4-5: 5-6: 4-5: 4-6: 5-6: 2, setae, long, acuminate
posteriorly. Sternal chaetotaxy: 0: 4: 4: 4-6: 4-6: 6: 4: 4: 4-6: 4-6: 6: 2. Sternites 3-5
divided. Coxal chaetotaxy: 3-4 coxal spines in medial position along anterior margin of
ZnCl, COs 22 Ze i, Ws Zevs OO Zeal, Os evs a, Oe ale

Dimensions (mm): body length 0.48-0.63; pedipalp: trochanter 0.06-0.07/0.05,
femur 0.12-0.16/0.04-0.07, tibia 0.07/0.05-0.07, chela (with pedicel) 0.25-0.28/0.06-0.07,
chela (without pedicel) 0.23-0.27/0.06-0.07, movable finger length 0.14-0.19; carapace
0.19-0.23/0.16-0.20.

Protonymph. L/W ratio: pedipalpal trochanter 1.25-1.33, femur 2.20-3.00, tibia
1.25-1.50, chela (with pedicel) 3.33-4.00, chela (without pedicel) 3.16-3.80 x longer than
broad. Fixed finger with 3 trichobothria, ist, et, eb present; movable finger with 1
trichobothrium, ¢ present; serrula exterior of chelicera with 7-8 broad, elongate
lamellae.

Carapace: anterior margin smooth, slightly dentate medially, 4: 2: 5: 3 setae, 1.06-
1.16 x longer than broad (ocular and posterior width equal). Tergal chaetotaxy: 2: 2-3: 2-
3; 2-3: 42 3: 3-4; 3: 4: 3-42 4-5: 2. Sternal chaetotaxy: 0; 2-323: 4: 2-3: 2-4: 4: 4: 3-4: 3-4:
4: 2. Coxal chaetotaxy: 3-4 small coxal spines along anterior margin of coxa 2, 2: 2: 1, 0:
2 (0. Os 2 ©, Os WE7s sips deal

Dimensions (mm): body length 0.36-0.45; pedipalp: trochanter 0.04-0.05/0.03-
0.04, femur 0.09-0.11/0.03-0.05, tibia 0.05-0.06/0.04, chela (with pedicel) 0.17-0.22/0.05-
0.06, chela (without pedicel) 0.16-0.21/0.05-0.06, movable finger length 0.11-0.14;
carapace 0.14-0.17/0.12-0.16.

DISCUSSION

Only two species of the genus Austrochthonius Chamberlin are recorded from
Australia namely: A. australis Hoff and A. cavicola Beier (1968). Results of this investiga-
tion are based on the examination of specimens from locations ranging from lat.
33°45'S, long. 150°57’E to lat., 41°49’S, long. 145°37’E and suggest a considerable
size variation exists within A. australis which is not consistent with clinal variation.

While size is important, body length was not considered a reliable guide in itself
because measurements can vary considerably due to a number of factors; for example,
the freshness of material, the inclusion of gravid specimens, the nature of preservative
and length of time in fixative. Consequently, the results of this research are based on
measurements of important specific morphological features which remain relatively
constant within populations.

A comprehensive description of the nymphal population is presented for the first
time and is therefore applicable only to the small specimens recovered from the
Cumberland State Forest, Sydney, N.SW. Lack of material representative of
southeastern Australia precluded a reliable assessment in this respect.

ACKNOWLEDGEMENTS

I wish to thank Dr. N. I. Platnick (American Museum of Natural History), Dr. M.
Gray (Australian Museum), Dr. M. S. Harvey (Western Australian Museum) for loan
of material and Dr. N. N. Tait for reviewing the manuscript.

References

BEIER, M., 1966 — On the Pseudoscorpionidea of Australia. Aust. J. Zool., 14: 265-283.
——, 1968 — Some cave dwelling Pseudoscorpionidea from Australia and New Caledonia. Rec. S. Aust. Mus.,
15: 757-758.

PROC. LINN. SOG. N.S.W., 111 (4), 1989

240 REDESCRIPTION OF PSEUDOSCORPION

CHAMBERLIN, J. C., 1923 — On two species of pseudoscorpion from Chile with a note on one from Sumatra.
Revista Chilena de Historia Natural, 27: 187-190.

——, 1929 — A synoptic classification of the false scorpions or chela-spinners, with a report on a cosmo-
politan collection of the same, Part I. The Heterosphyronida (Chthoniidae) (Arachnida — Chelon-
etida). Ann. Mag. nat. Hist., 4: 50-80.

, 1931 — The arachnid order Chelonethida. Stanford Univ. Publs. (Biol. ), 7: 1-284.

HANSEN, H. J., 1893. — Organs and characters in different orders of Arachnids. Ent. Medd., 4: 232.

Horr, C. C., 1951. — New species and records of Chthoniid pseudoscorpions. Am. Mus. Novit., 1483: 1-13.

LeGcG, G., 1975. — The genitalia and associated glands of five British species belonging to the family

Chthoniidae (Pseudoscorpiones: Arachnida). J. Zool. London, 177: 99-121.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

Pied Currawongs (Strepera graculina): their Diet
and Role in Weed Dispersal in suburban Sydney,
New South Wales

ROBIN A. BUCHANAN

BUCHANAN, R. A. Pied Currawongs (Strepera graculina): their diet and role in weed
dispersal in suburban Sydney, New South Wales. Proc. Linn. Soc. N.S.W. 111 (4),
1989: 241-255.

In this dietary study of an abundant population of pied currawongs (Strepera
graculina) 1009 regurgitated pellets, collected over 25 months, were analysed. The
percentage of pellets containing plant parts, mostly seeds of fleshy fruit, was high
throughout the year (79-98%). The fruit of introduced plants was present in 45-91% of
pellets containing plant material. The seeds of 46 species, of which 36 were introduced,
were identified in their pellets.

Fruits of the family Oleaceae, including the three introduced species, Ligustrum
sinense (small-leaved privet), Ligustrum lucidum (large-leaved privet) and Olea africana
(wild olive) were the most significant part of the currawong diet for three months of the
year, when at least one of these species was present in 54-74% of the pellets analysed.
Other major plant species in the diet included the introduced Pyracantha angustifolia,
Morus nigra, Ochna atropurpurea, Solanum pseudocapsicum, and the native Elaeocarpus
reticulatus.

The proportion of pellets containing animal parts decreased from a value of 50-
75% in the warmer months to 11-12% in the coldest months of the year. Few vertebrate
remains were found in the pellets; the major animal components identified were bull-
ants (Myrmecia spp.), beetles, and other insects.

Robin A. Buchanan, 22 Alicia Road, Mt. Kuring-gat, Australia 2080; manuscript received 22
December 1987, accepted for publication in revised form 19 April 1989.

INTRODUCTION

A varied diet, intelligence and bold behaviour have contributed to making the pied
currawong (Strepera graculina) one of the most successful and abundant birds in suburban
environments throughout eastern Australia (Readshaw, 1968; Wimbush, 1969; Blakers
et al., 1984). Within twenty years of European colonization, pied currawongs had
exploited new habitats and foods provided by settlement (Currey, 1966); and by the
1920’s they were listed as one of the main dispersal agents of Opuntia spp. (Anon, 1920;
1927). More recently it has become clear that they are also agents for the dispersal of two
abundant privet species (Ligustrum sinense, L. lucidum) in suburban Sydney (Walsh, 1965;
Vellenga, 1966; Rose, 1973; Clyne, 1980). Despite their large size (41-51cm), abundance
and role in weed dispersal, the diet of pied currawongs has not been studied in detail.

Rose (1973) systematically sampled regurgitated pellets, but many other accounts
are largely anecdotal. Reports of their animal prey occur in Lea and Grey (1936),
Marshall (1935), Roberts (1942), Readshaw (1965), Recher (1976), and Cooper and
Cooper (1981) and include many insect orders as well as worms, snails, crabs, birds,
mammals and carrion. Items of fruit eaten are frequently listed in catalogues of regional
birdlife (Table 1).

From October 1976 to October 1978 the pellets regurgitated by pied currawongs in
a garden in suburban Sydney were collected and their contents analysed. Objectives of
this survey were: to record details of the larger contents of pellets, relating them to
seasonal abundance of dietary components; to discuss the dietary findings in relation to
the role of pied currawongs in weed dispersal in the Sydney region.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

242

CURRAWONGS AND WEED DISPERSAL

TABLE 1

Fruits and seeds reported in previous studies and additional to those recorded from Thornleigh

Identification*

MONOCOTYLEDONES

Poaceae

*Maize
DICOTYLEDONES
Anacardiaceae

*Schinus molle

*Pepper Tree

*Pepper Tree
Aquifoliaceae

*Holly Tree
Cactaceae

*Prickly Pear

*Prickly Pear
Loranthaceae

Mistletoe

Loranthus

Mistletoe

Mistletoe
Malvaceae

#Lagunaria patersonia
Mimosaceae

# Paraserianthes lophantha
Moraceae

Moreton Bay Fig

Ficus macrophylla
Myrtaceae

#Eucalyptus caesia
Philesiaceae
Rosaceae

*Loquat

*Wild Raspberry

*Rowan

*English Laurel
Rutaceae

*Orange and Lemon
Santalaceae

Exocarpus stricta
Solanaceae

*Chillies
Vitaceae

Native Grape

x =

* = introduced to Australia.

# =

+ N = notstated.
O = observation while feeding.
P = pellet
S = stomach content.

Author

Edwards 1922

Rose 1973

Cheney 1915

Bourke 1949

Vallenga 1966

Morse 1922

Anon 1920

Anon 1927

Cleland, Maiden, Ferguson & Musson 1918
Lea and Gray 1936

Keast 1958

Rose 1973

Smith, Larkins & Pegler 1984
Smith, Larkins & Pegler 1984

Agnew 1922
Robertson 1969

Smith, Larkins & Pegler 1984
Rose 1973

Roberts 1942
Marshall 1935
Vallenga 1966
Vallenga 1966
Roberts 1942
Lea & Gray 1936
Bravery 1970

Cleland, Maiden, Ferguson & Musson 1918

indigenous to Australia but not to the Thornleigh area.

SITE DESCRIPTION

Type of Record +

QO OOOZdZ

The common or scientific name is listed in the same form as that given in the relevant paper.

The study was carried out at Thornleigh, 19km north-west of Sydney, New South
Wales; area features are illustrated in Fig. 1. Thornleigh is situated on a ridge 1-3km
wide with the steep-sided valleys of the Lane Cove River to the south-east and Berowra
Creek to the north-west. Both valleys are forested and the land to the west of Berowra

PROC. LINN. SOC. N.S.W., 111 (4), 1989

R.A. BUCHANAN 243

Creek is rural. Most of the suburban development consists of detached dwellings in
gardens well vegetated with native and introduced trees and shrubs. The percentage of
suburban development within a one to three kilometre radius of the study site ranges
from 50 to 67% respectively, with the remainder being natural vegetation.

E D railway

highway ~

built-up

Collection site @

Fig. 1. Map of the environs of the collection site showing the distribution of the built-up areas and native
vegetation.

The majority of the natural vegetation is woodland/open-forest (Specht, 1970) and
is included in the Sandstone Complex, both wet and dry sclerophyll, of Specht e¢ al.,
(1974). Most of the slopes are free of weeds but L. sznense and other weeds are abundant
along the creek and river banks.

Regurgitated pellets were collected at the junction of built-up and forested areas on
the eastern side of Thornleigh. The collection site was a lawn surrounded by a dense
growth of shrubs and trees; pied currawongs were attracted by bread, honey, seed, meat
and water.

PROG. LINN. SOG. N.S.W., 111 (4), 1989

244 CURRAWONGS AND WEED DISPERSAL

METHODS

Pellets

Pellets were collected twice a week over an area of approximately 150 m? from
October 1976 to October 1978. In total 1 009 pellets were examined; the highest number
sampled in any one month was 53 and the lowest 8 (Table 2). In October 1978 some 41
pellets were analysed.

TABLE 2
Number of pellets analysed in the 24 month period (October 1976-September 1978) and collected in the second year

2 48 4] 2
(Analysed)

2 119 64 24 2
(Collected)

The sampled pellets were divided into three classes of coherence; coherent
(collected intact or almost intact), partly coherent (collected in more than one piece),
not coherent (collected completely fragmented). The lengths and widths of coherent
pellets were measured and the contents of coherent and partly coherent pellets were then
separated by stirring them in water and filtering. All pellets separated into individual
fragments with this treatment. Only the larger fragments, identifiable at 40x magnifi-
cation, were noted.

Component Identification

Most seeds were identified by comparison with seeds from known plants and by
identification of young plants grown from seeds in the pellets. Some of the rarer seeds in
the pellets were identified by the National Herbarium (Sydney). Scientific and vernacu-
lar names are listed in Table 3; botanical and common names follow the National
Herbarium of New South Wales. Classification of invertebrates — based on Common-
wealth Scientific and Industrial Research Organization, 1970 — was only attempted to
class or order and rarely to genus. Many tiny invertebrate fragments remained un-
identified. Some bone fragments were identified by the Australian Museum, Sydney.
The number of different components, identified to class or genus for insects, and species
for plants was tallied for each pellet.

The plant species most frequently present in pellets was assessed by counting the
number of pellets in which a species was present in each month.

Bull-ants (Myrmecia species)

As the number of pellets containing bull-ants showed a strong seasonal pattern, an
estimate of the activity of bull-ants was obtained by observing six nests for 5-10 minutes
on one afternoon a week during the second year of the study. The largest number of ants
present on the surface of the nest during this time was recorded. The average number of
observed bull-ants was then calculated for each month.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

R. A. BUCHANAN 245

TABLE 3
Species of fruit recorded in pellets

Highest frequency in any one month (7%)

Classification (Common Name) 00. | 10-30 30-50 50-70 70-90

GYMNOSPERMAE
Cupressaceae

* Juniperus sp. (Juniper) x
MONOCOTYLEDONES
Arecaceae

*Phoenix sp. (Palm) x
Asparagaceae

#0* Protasparagus aethiopicus (Asparagus Fern)
Asteliaceae

+ Cordyline rubra

Phormiaceae

Dianella caerulea (Flax Lily)
Smilacaceae

Smilax glyciphylla (Thornless Smilax)
Zingiberaceae

#0* Hedychium gardnerianum (Ginger Lily)
DICOTYLEDONES
Anacardiaceae

#0* Toxicodendron succedaneum (Rhus) x
Araliaceae

0* Hedera helix (Ivy) x

Polyscias sambucifolia x
Cornaceae

* Dendrobenthamia capitata x
Ebenaceae

*Diospyros kaki (Persimmon) x
Elaeocarpaceae

Elaeocarpus reticulatus (Blueberry Ash) x
Euphorbiaceae

Omalanthus populifolius (Poplar-leaved Omalanthus) x
Lauraceae

#0* Cinnamomum camphora (Camphor Laurel) Xx
Magnoliaceae

0* Magnolia grandiflora (Evergreen Magnolia) x
Malvaceae

+ Lagunarta patersonia (Norfolk Island Hibiscus) x
Meliaceae

0+ Melia azedarach (White Cedar) x
Moraceae

* Ficus carica (Commercial Fig) Xx

Ficus rubiginosa (Port Jackson Fig) x

0* Morus alba (Mulberry) xX
Myrtaceae

Eucalyptus resinifera (Red Mahogany) x

+ Syzygium paniculatum (Lilly Pilly) x
Ochnaceae

#0* Ochna atropurpurea (Ochna) x
Oleaceae

#0* Ligustrum lucidum (Large-leaved Privet) x

#0* Ligustrum sinense (Small-leaved Privet) Xx

#0* Olea africana (Wild Olive) Xx
Phytolaccaceae

#0* Phytolacca octandra (Ink Weed) x
Pittosporaceae

Pittosporum undulatum (Pittosporum) x

x we Km KK

PROC. LINN. SOC. N.S.W., 111 (4), 1989

246 CURRAWONGS AND WEED DISPERSAL

TABLE 3 (Cont’d.)

Highest frequency in any one month (%)

Classification (Common Name)

Proteaceae

Persoonia pinifolia (Pine-leaf Geebung)
Rosaceae

#0* Cotoneaster glaucophyllus (Cotoneaster)

#0* Duchesnea indica (Wild Strawberry)

* Fragaria ananassa (Strawberry)

*Malus sp. (Apple)

* Prunus sp. (Plum and Cherry)

#0* Pyracantha angustifolia (Orange Firethorn)

*Pyrus sp. (Pear)

#0* Rhaphiolepis indica (Indian Hawthorn)

*Rubus x loganobaccus (Loganberry)
Rubiaceae

Morinda jasminoides
Rutaceae

* Citrus sinensis (Orange)
Solanaceae

#0* Solanum pseudocapsicum (Madeira Winter Cherry)
Sterculiaceae

0+ Brachychiton acerifolius (Iawarra Flame Tree)
Verbenaceae

#0* Lantana camara (Lantana)
Vitaceae

0* Parthenocissus sp. (Virginia Creeper)

* Vitis labrusca (Grape)

* = introduced to Australia

+ = indigenous to Australia but not to the Thornleigh area
0 = present in the forest

# = reproductive in the bushland

No notation = native to the area

RESULTS

Pellets

Occurrence

The smallest number of pellets collected occurred in the autumn months (Table 2),
a period which coincided with a seasonal decrease in the number of birds visiting the site
(Buchanan 1983). Only eight pellets were collected in April, a deposition rate of 0.05 per
square metre per month. The highest number occurred in September when 132 were
collected, a deposition rate of 0.9 per square metre per month. The decrease in the
number of pellets collected in July cannot be explained.

The average width of coherent pellets was 15mm with a standard deviation of 0.32.
The average length of coherent pellets was 27mm with a standard deviation of 0.64. The
maximum number of non-coherent pellets (Fig. 3) was recovered over the late autumn
and winter months of April, May, June, July and August, when the pellets lacked
animal remains to bind the seeds together (Fig. 4). The maximum number of coherent
pellets was recorded in October (58%) and November (71%) when pellets contained
large amounts of mulberry (Morus alba). The majority of these multiple fruits were
relatively intact.

The data collected over 25 months showed that feeding is concentrated on one or a
few items during the time needed to produce a pellet (Fig. 2). Almost 40% of pellets
contained one component and more than 70% contained one or two.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

R. A. BUCHANAN

30.

20.
pellets

3 4
NO components - 2 yrs_ data

Deal

Fig. 2. Percentages of pellets containing different numbers of different components for the 25 month period.

pellets

PARTLY COHERENT

Fig. 3. Average monthly percentages of coherent, partly coherent and non-coherent pellets.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

248 CURRAWONGS AND WEED DISPERSAL

100.
O O Q
O
7 a Q
O
® Oy H
80. 0 e O
C)
‘a } Wi,
rh es /
Daa! LC] /
D
60 ‘
%
CL]
Pellets 4
.

O re) plant
20, @——® animal

O—y introduced plants

month

Fig. 4. Average monthly percentages of pellets containing plant parts, introduced plants and animal parts for
the 25 month period.

Components and Seasons

Plant Content

The percentage of pellets containing plant parts remained high (79-98%) through-
out the two years (Fig. 4). Fruit and seed was the major component (Table 3) but leaves
and flowers were recorded in 3% and 0.6% of pellets examined. Almost half (22) of the
46 recorded fruits and seeds were present in less than 10% of the pellets in any one
month (Table 3). Twenty-one species were abundantly represented, (Fig. 5).

Oleaceae, including Ligustrum sinense, L. lucidum and Olea africana were present in
over 20% of pellets from May to September and peaked at 74% in July. L. sznense was the
most abundant and was present in over 20% of pellets for three months of the year (Fig.
6). The consumption of Olea africana was highest in the early winter months but never
peaked above 20% and L. lucidum was the most important in the later part of winter
(26% in August).

Two other common weeds of urban native vegetation, Ochna atropurpurea and
Cinnamomum camphora, were a prominent part of the diet in summer and autumn respec-
tively. Insufficient sampling may have exaggerated the importance of Cinnamomum

PROG. LINN. SOC. N.S.W., 111 (4), 1989

R.A. BUCHANAN 249

camphora in April of the second year — only eight pellets were collected. Morus alba
dominated the diet in October and November and Pyracantha angustifolia was consumed
for several months of the year (Fig. 5).

4st year and year
% pellets

80 60 40 20 20 40 60 80

Oo
Elaeocarpus reticulatus |___———] S_— Ligustrum lucidum*

MM alban® het ode ee ESE IM alba
J [ Persoonia pinifolia | ————__F Ochna atropurpurea’

a Ochna atropurpurea*{ si Ss CO. atropurrpureae
Prunus sp..|_____—*|_ ~_| Solanum _pseudocapsicum’*

Malus sp."|__——d| ds Phenix sp.”
O. atropurpurea" |__| Vitis labrusca’

Lantana camara*

F | Ficus carica’____ ———————_—sS's«@Ficus carica®
Vitis labrusca® |_| SSCCSd’:séPhoolenix sp’

P. angustifolia”

Cinnamomum camphora® ee S. pseudocapsicum *

M Melia azedarach’
Syzigium oleosum*|__ Cd Ss. angustifolia”

J Ligustrum_sinense * Ligustrum_sinense*
O._africana* |___——*d|s«~Pittosporum undulatum

J
Llucidum’ [Ys ucidum*

L. lucidum * |_| L. sinense’

| L.sinense’ | CSC. cidum
A
|____J} P angustifolia®

E. reticulatus |__| Elaeocarpus reticulatus

L. sinense* |___|_| L. sinense’
|__| L. lucidum*
|__| Smilax glyciphylla

Fig. 5. The two plant species most frequently present in pellets for each month, showing changes between the
first and second years of the 25 month period.

Introduced plants were present in 50-90% of pellets in all but one month (Fig. 4).
Twenty-one of the 36 introduced plants recorded are present in the bushland and at least
15 of these produce seed in this situation. Only 10 of the 46 species were native to the
Thornleigh area. The most important of these were Elaeocarpus reticulatus, Persoonia pint-
folia, Pittosporum undulatum and Smilax glyciphylla (Fig. 5). Only Elaeocarpus reticulatus was

PROC. LINN. SOC. N.S.W., 111 (4), 1989

250 CURRAWONGS AND WEED DISPERSAL

the dominant native species in any month (September). Prttosporum undulatum, which has
a similar habit and similar habitat requirements to Ligustrum sinense and which is
dispersed over the same months as the Oleaceae, was sparingly eaten.

80,

w---@ Oleaceae w Y

60.
oO—o Ligustrum sinense

% e @ Olea africana

pellets oO — o Ligustrum lucidum

40.

20.

month

Fig. 6. Average monthly percentages of pellets containing Oleaceae, Ligustrum sinense, Ligustrum lucidum and
Olea africana.

The data in Table 4 indicate that the fruit of the most favoured species have an
average diameter greater than 5mm. Fruits of several species (Morus alba, Ficus sp.,
Solanum pseudocapsicum and Pittosporum undulatum) contained many seeds; but, the
majority of fruit eaten contained only one or two seeds. For this latter group, the smaller
the fruit is, the higher the number of seeds contained in a pellet. Fruits 5-6mm in
diameter were represented by an average of 8-42 seeds per pellet while large fruit such as
Persoonia pinifolia (14mm), Syzygium paniculatum (14mm) and Melia azedarach (13mm) were
only represented by 2-5 seeds per pellet. Flesh and skin, but no seeds, of the very large
fruited apple and persimmon were recorded in the pellets. The maximum number of
seeds per pellet for Ligustrum sinense was 162, Lantana camara 131, Ligustrum lucidum 67,
and Ochna atropurpurea 42.

The major fruits in the currawong diet ranged in colour from bright orange
(Diospyros kaki, Pyracantha angustifolia and Solanum pseudocapsicum) to green (Vits labrusca
and Persoonia pinifolia) to black-blue for 7 of the 21 most frequent species.

Fruit Selection

Only ripe fruit was eaten or ingested by the pied currawongs. For green and pale-
coloured fruit pied currawongs appeared to use means other than colour to determine
ripeness. Just before Melia azedarach fruit first appeared in pellets, individual currawongs
were observed to visit the tree; in each case the bird selected several fruits, held each one
in the beak for a few seconds, and then dropped it. It is suggested that the birds were

PROC. LINN. SOC. N.S.W., 111 (4), 1989

R. A. BUCHANAN 251

assessing the yellowish coloured fruit for ripeness by testing the softness of the pericarp
with their beaks or tasting the flavour of exuded juices.

TABLE 4

Fruit size and colour, number of seeds per fruit, average and maximum number of seeds in pellets

Species

Malus domestica

Diospyros kaki fruit very large — only skin and flesh ingested, no seed

may have many hundreds of seeds per pellet

Phoenix sp. 1 3
Ficus sp. may have many hundreds of seeds per pellet

Prunus nigra approx 1 1

20
Pittosporum orange only approx approx
undulatum consume 50 200

seed

Vitis labrusca

Persoonia pinifolia

Syzygium
paniculatum

Melia azedarach

Solanum orange approx approx
pseudocapsicum 60 300

Cinnamomum black 4 23
camphora

Elaeocarpus reticulatus blue 22

Pyracantha orange approx
angustifolia 60

Olea africana 8
Ligustrum sinense 6 36
Ligustrum lucidum 6 18 67
Lantana camara 5 131

Smilax glyciphylla 3) 54

PROC. LINN. SOC. N.S.W., 111 (4), 1989

252 CURRAWONGS AND WEED DISPERSAL

Animal Content

The percentage of pellets containing animal parts was highest (50-75%) in the
warmer months but decreased to a low of 11-12% in the two coldest months of the year,
June and July (Fig. 4). Remains of 7 major animal groups were recorded; these were
Gastropoda, Arachnida, Diplopoda, Insecta, Amphibia, Aves and Mammalia.

Vertebrate remains were rare in the collected pellets and only 14 contained bone
fragments (Table 5). The fragments in seven of the pellets were too small for identifica-
tion, but of the remaining seven, two were from birds, two from frogs, two from house
mice (Mus musculus) and one was a chicken bone.

The majority of the animals eaten were invertebrates — especially insects. Adult
beetles, lepidopteran larvae and hymenopterans, — particularly bull-ants — were the
most abundant remains (Table 5). The percentage of pellets containing bull-ants was
high in the summer months, peaking at 47% in December, but fell to 0-4% during the
winter months. This decrease was correlated with reduced bull-ant activity outside the
nest (Fig. 7).

TABLE 5

Animal remains identified in pellets

Month and Number of Pellets Containing Item

Categories of Animal Remains ), SES IMO VAS MiG]! AG SS = O= SNe
Gastropoda (snails) 2 Z
Arachnida (spiders) 1
Diplopoda (millipedes) 1
Insecta
Ephemeroptcra (mayflies) 1
Blattodea (cockroaches) 1 1
Mantodea (praying mantids) 1
Dermaptcra (carwigs) 3 1
Orthoptera (grasshoppers & crickcts) 1 ar 1 1 2 1
Hemiptera (bugs) 2
Coleoptera (adult beetles) 30 16 9 8 10 3S & F 4 Ol BW sé
Lepidoptera (moth & butterfly larvac) Daa! alle le ei ec sais) = 2 OR Soon |)
other 1 7 saa \ ae ae 1 2,
Hymenoptera (sawflies, wasps, bees & ants)
Myrmecia sp. (bull-ant) 223 Oe ee On 3.20 52 26 40
other SOY el Se a TA ett aaa) 1 SIE 93 3 I @
Unidentified 0 12° 10 @ 8 22 lily te e20) (25s
Bone Zar RMe E7 A alk ihre ae 2 2 1
Egg shell i 2

Number of pellets examined Y @7 42 47 GW) SY D7 BY GH 140 Br BH

* = 3 months’ data

DISCUSSION

Pellets and Diet

The examination of regurgitated pellets does not reflect the complete diet since
only indigestible items are recorded (Dorst, 1971). The comparative volume of items in
the pellets is also only a general guide to their importance in the bird’s diet. For example,
remains of lepidopteran larvae were usually only represented by the head capsule even
though the larvae may have contributed a significant portion of the day’s intake of food.
On the other hand, favoured fruit items such as Elaeocarpus reticulatus, Olea africana, Ligus-
trum sinense and L. lucidum have thin fleshy layers around a hard endocarp and lost very
little of their volume after digestion.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

R. A. BUCHANAN DDS

Despite the drawbacks outlined above the pellet studies reported here have aug-
mented information on currawong diets in several ways. The importance of plant
material in the omnivorous diet has been confirmed, but the wide diversity of fruits and
seeds ingested — including many introduced species — has been documented in detail;
the variety of animals eaten has also been demonstrated. Seasonal frequency in a
number of animal and plant remains, and the small number of dietary components in
individual pellets constitutes further evidence that currawongs are opportunistic
feeders.

Myrmecia sp.

bull-ants

pellets

J
month

Fig. 7. Percentages of pellets containing bull-ants (Myrmecza spp.) in the 25 month period and the average
number of bull-ants recorded at nest surfaces for one year.

Fruit Selection and Dispersal

The utilization of a large number of introduced plants as food resources by the
currawong has wider implications, because of the potential for dispersal of weed species.
Dispersal is only possible if the seed is undamaged by ingestion. Many seeds included in
the pied currawong pellets were identified by germinating them and viability was high.
Ligustrum sinense viability ranged from 83-91% (three samples of 50, 46 and 12 seeds),
Pittosporum undulatum 100% (8 seeds) and Toxicodendron succadaneum 100% (6 seeds).

The Oleaceae (Ligustrum sinense, L. lucidum and Olea africana) seem to be ideally
suited to dispersal by pied currawongs. The prolific crop is produced in winter at a time
when pied currawongs form large feeding flocks in the warmer parts of their range,
including Sydney (Buchanan, 1983). The black-blue colour of the fruit is one of the
colours preferred by birds (Turcek, 1963). The size of the fruit is at the lower limit of the
size taken by pied currawongs so that large numbers of fruit, and hence seeds are con-
sumed. The ratio of fruit to animals eaten also increases in the winter months so that a
higher proportion of fruit may be eaten than in summer. Deposition of at least some of
the seed in suitable conditions for germination and establishment (i.e. along creeks and

PROG. LINN. SOC. N.S.W., 111 (4), 1989

254 CURRAWONGS AND WEED DISPERSAL

rivers) 1s ensured by the fact that pied currawongs usually regurgitate pellets after
drinking (Robertson, 1969; Clyne, 1980).

The average number of seeds deposited per square metre can be large; in June 1977
approximately 12 Ligustrum sinense seeds were dropped on every square metre of the
collection site. The density of deposition away from the collection site would be much
less, but the common occurrence of clustered Ligustrum sinense, L. lucidum and Cinna-
momum camphora seedlings in natural vegetation confirms that seed dispersal by birds is
important.

Some fruit appeared to be rejected by the birds. For example, Cotoneaster
glaucophyllus is abundant in the study area and carried a prolific crop of fruit each
autumn and winter, but was never present in more than 10% of pellets in any one month
(Table 3). Rejection of Cotoneaster glaucophyllus cannot be explained. It was not on the
basis of the thickness of the flesh (1.4mm) as this was greater than for Elacocarpus reticu-
latus and the Oleaceae (0.8-1.2mm). Fruit size was within the range of the most
frequently eaten species (Table 4) and by the colour it should have been selected (Turcek
1963).

As well as the dispersal of obviously palatable items, pied currawongs may also
distribute the seed of woody fruit. The fruit of Eucalyptus resinifera was found in one
pellet. Even if the inclusion of such fruit and viable seeds 1s a rare event, the possibility
for dispersal of minor dietary items is important.

ACKNOWLEDGEMENTS

My thanks are due to S. Buchanan for helping to collect the pellets and for the
typing. The National Herbarium (Sydney) and the Australian Museum identified some
of the more difficult items. My thanks are also due to H. Recher who commented on the
manuscript.

References

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RECHER, H. F., 1976. — Reproductive behaviour of a pair of pied currawongs. Emu 76: 224-226.

Roserts, N. L., 1942. — The winter flocking of the pied currawong. Emu 42: 17-24.

ROBERTSON, J. S., 1969. — Measurements, weights and notes on pied currawongs. Aust. Bird Bander 7: 7-10.

Rose, A. B., 1973. — Food of some Australian birds. Emu 73: 177-183.

SMITH, L., LARKINGS, D. and PEGLER, J., 1984. — Garden plants attractive to birds. Aust. Birds 18: 17-27.

SPECHT, R. L., 1970. — Vegetation. Jn LEEPER, G. W. (ed.) The Australian Environment. Melbourne:
C.S.I.R.O., Melb. Univ. Press.

SPECHT, R. L., ROE, E. M. and BOUGHTON, V. H. (ed.), 1974. — Conservation of major plant communities
in Australia and Papua New Guinea. Aust. J. Bot. Supplement No. 7.

TURCEK, F. J., 1963. — Colour preference in fruit and seed-eating birds. Proc. XIII Intern. Ornithol. Congr:
285-292.

VELLENGA, R. E., 1966. — Notes on the pied currawong. Aust. Bird Bander 4: 6.

WALSH, J. E., 1965. — Notes on the pied currawong. Aust. Bird Bander 3: 53.

WIMBUSH, D. J., 1969. — Studies on the pied currawong Strepera graculina in the Snowy Mountains. Emu 69:
72-80.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

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Nine New Deep-water Species of Echinodermata
from Norfolk Island and Wanganella Bank,
northeastern [asman Sea, with a Checklist of the
Echinoderm Fauna

FRANCIS W. E. ROWE

Rowe, F. W. E. Nine new, deep-water species of Echinodermata from Norfolk Island
and Wanganella Bank, northeastern Tasman Sea, with a checklist of the echino-
derm fauna. Proc. Linn. Soc. N.S.W. 111 (4), 1989: 257-291.

Nine new species comprising 3 holothurians, 2 echinoids, 3 asteroids and 1
crinoid are described from the deep waters surrounding Norfolk Island and Wanganella
Bank, northeastern Tasman Sea. A checklist of the echinoderm fauna shows at least 123
species are now known for that region, which includes the nine new taxa and seven taxa
identified only to the level of genus. Taxonomic notes are included in the checklist for
the taxa Bathyplotes punctatus (Sluiter); Holopneustes inflatus Lutken in Agassiz; Ophiothrix
(Acanthophiothrix) lepidus de Loriol; Fromia polypora H. L. Clark; Coscinastertas muricata
Verrill; Astrostole rodolphi (Perrier); Oxycomanthus plectrophorum (H. L. Clark) and Cenolia
spanoschistum (H. L. Clark).

Francis W. E. Rowe, Division of Invertebrate Zoology, Australian Museum, PO. Box A285, Sydney
South, Australia 2000; manuscript received 7 December 1988, accepted for publication 19 July
1989.

INTRODUCTION

Norfolk Island (29°S, 168°E) and Wanganella Bank (c. 32°32’S, 167°32’E) lie to
the northwest of the northwest tip of New Zealand. A number of recent, published
accounts describe, or include records of echinoderms from those regions (Pawson,
1965b; McKnight, 1967, 1968a,b, 1975, 1977; H. E. S. Clark, 1970, 1982; Baker, 1979,
1980; Rowe, 1977, 1985; Edgecombe and Bennett, 1983; Rowe and Albertson, 1987).
During the course of an investigation of the systematic composition and zoogeographic
relationships of the echinoderm fauna of New South Wales (Australia), Lord Howe
Island and Norfolk Island (Tasman Sea), the author has examined echinoderm material
held in the collections of the Australian Museum (AM), Museum of Victoria (MV),
National Museum of New Zealand (NMNZ) and the collections of the New Zealand
Oceanographic Institute (NZOI). This latter collection includes, particularly, a great
deal of material from Norfolk Island and Wanganella Bank along the Norfolk Ridge.
Previous publications have recorded nearly 70 species from those regions. The present
study has revealed not only nearly double that number (123 species) can be recorded,
but that 9 of those are species which are new to science and seven taxa are identified as
far as genus. The purpose of this paper is to provide descriptions of the new species and
present a preliminary, updated checklist of all the taxa now recognized. The checklist
includes a citation of the reference recording the taxon from the vicinity of Norfolk
Island or Wanganella Bank; a general distribution similarly with an appropriate refer-
ence and/or indication to ‘this work’ indicating the present author’s view; depth range;
brief taxonomic note for each of 8 species where a comment is required. New records
from Norfolk Island or Wanganella Bank are denoted by an asterisk (*). For the pur-
poses of this report an area delimited by the latitudes 28°-33°S and longitudes 167° -
169°E is used to include records from waters surrounding both Norfolk Island (28° -
30°S, 167° -169°E) and Wanganella Bank (>30°-33°S, 167° -169°E).

PROG. LINN. SOC. N.S.W,, 111 (4), 1989

258 NEW TASMAN SEA ECHINODERMS

SYSTEMATIC ACCOUNT
Class HOLOTHURIOIDEA
Family Holothuridae

Holothurta ( Vaneyothuria) uncia n. sp.
Fig. 1A-D

Diagnosis: A species in the subgenus Vaneyothuria which is distinctively coloured uni-
formly cream with a chocolate brown ring around each tube-foot and with the rim of the
disc of the 3-dimensional table spicules smooth.
Material examined: Holotype, AM J21696, 29°24.8’S, 168°10’E, off Norfolk Island,
342-360 m (NZOI stn 191).
Description: The contracted holotype is about 135mm long and about 57mm wide at
the middle of the body. There are 17 tentacles. The tube feet are in three bands along the
ventral ambulacra. The ventral-lateral bands are formed by a zigzag alignment of the
pairs of tube feet. The mid-ventral band comprises two discrete rows of paired tube feet,
the rows spaced about 6-8mm apart. Dorsally the tube feet are more or less scattered,
without linear arrangement. The body wall is relatively thin. The calcareous ring is
stout and not unusual (Fig. 1A). Tentacle ampullae are present. The gonad comprises a
large bunch of branched and unbranched tubules on the left hand side of the dorsal
mesentery. It lies about 20mm from the anterior end of the body, and the tubules are up
to 60mm long. The respiratory trees are not unusual. The gut is full of sand.

Spicules of the tentacles comprise curved spinous rods. The largest rods, from the
tentacle stalk, measure up to about 480 wm long x about 50 wm wide. The smallest rods,
from the tentacle branches, measure 50-150 wm x 5-7.5 ym (Fig. 1B).

Spicules from the dorsal and ventral body walls comprise tables and buttons. The
disc of the tables are squarish to irregularly rounded, smooth-rimmed, 90-105 wm in
diameter and with a single ring of 8-10 holes (Fig. 1C). The spire comprises four pillars,
joined by one or two cross-beams. The crown of the spire is small (c. 20-25 pm
diameter), with a variable number of small spines. The spire is between 70-90 ym in
height. The buttons are usually smooth, irregular in outline and usually have three pairs
of holes (Fig. 1D). Rarely one or two low knobs may occur along the midline. They
measure between 52 wm long x 37 wm wide to 127 wm long x 97 wm wide, with the
majority measuring 90 wm x 52 pm. Tables in the dorsal tube feet range up to 112 pm disc
diameter, with up to 16 small, peripheral holes, either in a single ring or with some holes
offset into a partial, second ring. Buttons are present, but in addition curved and
straight supporting rods occur. An irregular end-plate is also present, c. 180 pm
diameter. Spicules in the ventral tube feet are often of similar type but the tables range
in size from disc diameter 50-90 um and spires from 40-55 wm in height. The buttons are
often elongate (165 wm long x 52 wm wide), with up to 6 pairs of holes. Supporting rods
are similar to those in the dorsal appendages and the end-plates range up to 400 wm
diameter.

Colour: Uniformly cream, with a narrow basal ring around the ventral tube feet and a
much broader (3-4mm diameter) ring of chococolate brown around the dorsal tube feet.
Etymology: Named for Uncza, the snow-leopard.

Remarks: This species (uncia) is clearly related to H. (V.) integra Koehler and Vaney, of
which H. neozaelanica Mortensen is a synonym (Rowe, 1969; Cherbonnier and Feral,
1981) and which is distributed from the Bay of Bengal, off Port Hedland, northwest
Australia, the Philippines and New Zealand. The spiny-disced tables of integra easily
separate the two species. Also, the colour of wncza is distinctive among known deeper-
water species of Holothuria.

PROC. LINN. SOG. N.S.W., 111 (4), 1989

FW. E. ROWE 259

Fig. 1. A-D Holothuria (Vaneyothuria) uncia n. sp. (holotype AMJ21696), A = calcareous ring, interradial and
radial plate, B = rods from tentacle, C = tables from body wall, D = buttons from body wall; E-H Mesothuria
(Penichrothuria) norfolkensis n. sp. (holotype AMJ21697), E = calcareous ring, F = rods from tentacle, G =
tables from body wall, H = ‘reduced’ table from tube-foot; I-M Neothyonidium parvipedum n. sp. (holotype,
AMJ21698), I = lateral view of left side of holotype, J = calcareous ring, K = rods from tentacle, L = tables
from body wall, M = end plate from tube foot.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

260 NEW TASMAN SEA ECHINODERMS
Family Synallactidae

Mesothuria (Penichrothuria) norfolkensis n. sp.
Fig. 1E-H

Diagnosis: A species in the subgenus Penichrothuria which is uniformly creamy-grey in
colour; tube feet are scattered over the body but most prominent along the ventral-
lateral ambulacra; reduced tables few; normal tables with 4 pillars which are crowned
with ring of numerous small spines.

Material examined: Holotype, AM J21697, 28°57.9’S, 167°45.5’E, off Norfolk
Island, 392-423m (NZOI stn P35).

Description: The holotype measures about 165mm in length and about 67.5mm in
width. It has 17 tentacles. The tube feet are scattered over the body, but are most promi-
nent along the ventral-lateral ambulacra.

There are no tentacle ampullae. The polian vesicle is single. The madreporite lies
to the right hand side of the dorsal mesentery. The gonad is a single tuft on the left hand
side of the dorsal mesentery. It comprises some 25 tubules, each ending in a number of
bifid lobes giving the gonad the appearance of a bunch of grapes. The water vascular
ring 1s separated from the posterior end of the calcareous ring by about 15mm. The gut
is packed with fine sand. The calcareous ring is stout, with radial plates deeply notched
posteriorly (Fig. 1E).

Spicules from the tentacles comprise more or less straight, or slightly curved, spiny
rods. The largest, from the stem, range in length and width up to about 400 ym x 75 pm,
respectively. The smallest rods occur in the branches of the tentacles and range between
about 60-120 wm in length (Fig. 1F).

Spicules from the body wall comprise tables only. Supporting rods and perforated
disc plates are additionally present in the tube feet. Rarely in the tube feet are there
small, reduced tables. The tables of the dorsal body wall have a disc diameter of 65-130
pm (Fig. 1G). There is either a single peripheral ring of 8-10 holes or an inner ring of 8
larger holes alternating with a partial or complete outer ring of 4-10 holes. The spire
comprises four pillars with one cross beam. The spires are 52-75 wm high and crowned
with a ring of numerous small spines. The tables of the ventral body wall are slightly
smaller, with disc diameter up to about 112 um. The tables of the dorsal tube feet are
smaller with disc diameters ranging from 50-90 um and spires up to 52 wm high. Other-
wise, the tables have similar appearance to those of the body wall. The supporting rods
are curved, perforated terminally and either side centrally. They are up to 187 um long.
The end plates are up to 300 wm in diameter. Small, irregular, reduced tables are
present, but very few in number (Fig. 1H). Spicules of the ventral and ventral-lateral
tube feet are similar, in all respects, to those of the dorsal tube feet, except that the end
plate of the ventral-lateral tube feet have a diameter up to 550 pm.

Colour: Uniformly creamy-grey.

Etymology: Named for the locality of discovery, Norfolk Island.

Remarks: The form of the tables, with a complete ring of numerous small spines at the
apex of the spire easily separates this species from all others, including M. (P) verrilli
(Theel) and M. (P.) carnosa Fisher, in the subgenus Penichrothuria. Were it not for the
occurrence of ‘reduced’ tables, albeit few, in the tube feet, I would consider M. (P.)
norfolkensis closely related to M. (Allantis) intestinalis (Ascanius), though separable from it
on the shape of the normal tables (the spire is more slender in intestinalis). Heding (1942)
is quite emphatic that the absence of ‘reduced’ tables gives a clear limit to the subgenus
Allantis in which he placed intestinalis. Because of the apparent rarity of the ‘reduced’
tables in M. (P.) norfolkensis, a fresh assessment of this character would be usefully under-
taken. This is particularly relevant because, in their rarity, these tables can easily be

PROC. LINN. SOC. N.S.W,, 111 (4), 1989

F. W. E. ROWE 261

overlooked. It is on this somewhat shaky character alone that norfolkensis is subgeneri-
cally separated from its apparent nearest relative, intestinalis.

Family Phyllophoridae

Neothyonidium parvipedum n. sp.
Fig. 1I-M

Diagnosis: A species of Neothyonidium with a caudal process, small tube feet in double or
irregular quadruple rows on the ventral ambulacra, more or less scattered dorsally;
calcareous ring tubular, polyplacous, radials with very long posterior processes, inter-
radials almost as long as the processes of the radials; two-pillared tables in body wall.
Material examined: Holotype, AM J21698, 32°36.32°S, 167°30.7’E, Wanganella
Bank, 126m (NZOI stn P4).
Description: The contracted holotype is about 15.25mm long and about 6mm wide at
mid body. The body is bluntly tapered anteriorly, but narrowing to a distinct caudal
process (tail) posteriorly. The tail is dorsally directed, about 2.5mm from the ventral,
backward projection of the body (Fig. 11). The tube feet are small, in double rows in the
ambulacra anteriorly and posteriorly. Mid-ventrally the tube feet are widely spaced in
somewhat irregular rows with 4 rows of tube feet in each ambulacrum. Dorsally the
podia are fewer in number and without apparent regular arrangement.

There are 20 tentacles, 5 pairs of larger alternating with 5 pairs of small tentacles.
The calcareous ring is polyplacous, massive, tubular, about 13.25mm long. The radial
plates are notched anteriorly, and have very long posterior processes. The interradial
plates are pointed anteriorly and long posteriorly, extending to within about 3mm of the
end of the ring. The interradials do not have posterior processes (Fig. 1J). The polian
vesicle and madreporite are each single. The form of the gonad 1s difficult to determine
but appears branched. It is preserved in a fused condition.

Spicules of the tentacles comprise smooth, straight, curved, S-shaped or, rarely,
X-shaped rods (Fig. 1K). The rods are terminally expanded and perforated. They range
in size from 75 um long x 3.5 ym wide to 240 wm long x 17 wm wide.

Spicules in the introvert comprise two-pillared tables. The disc is irregularly oval to
squarish. It has 4 large central holes surrounded by up to about 26-30 smaller holes in
one or two alternate rings. The disc reaches a maximum diameter of 112 um. The spire
measures up to 75 wm in height, has a single cross-beam and each of the two terminally
divergent pillars bears 2-3 spines at the tip.

Spicules in the body wall comprise scattered, two-pillared tables (Fig. 1L). The disc
of the tables range from 90 wm x 75 pm to 112 wm x 105 pm. The disc comprises 4 large,
alternating with 4 small peripheral holes. Rarely, an outer, incomplete third row of 3-4
smaller holes is present. The spire comprises two pillars joined by a cross-beam. The tips
of the pillars diverge and each is usually minutely bifid. The spires range from 40-60 ym
in height.

There are only end plates in the tube feet and these range in diameter from 75 pm
(anteriorly and posteriorly) to 100 wm (mid-ventrally) (Fig. 1M).

Colour: Uniformly greyish-white.

Etymology: Named for its small tube feet.

Remarks: This species is most closely related to N. hawazense (Fisher). It differs from
hawauense in possessing a caudal process, in the slightly smaller disc plates from the tube
feet, in possessing rods in the tentacles, and in the smaller number of perforations in the
discs of the tables in the introvert. It is possible that parvipedum will be found to be
conspecific with hawazense, but insufficient material does not permit such a conclusion
at this time. I am not convinced by Heding and Panning’s (1954) synonymy of Fisher's

PROG. LINN. SOG. N.S.W., 111 (4), 1989

262 NEW TASMAN SEA ECHINODERMS

Fig. 2. Hapalosoma pulchrum n. sp. (holotype, AMJ21699), A = aboral view, B = oral view (h.d. = 67.4mm).

PROC. LINN. SOC. N.S.W,, 111 (4), 1989

F. W. E. ROWE 263

(1907) second Hawaiian species N. alexandri with hawazense. There appears to be as close
a relationship between alexandri and N. armatum Pawson, 1965c, from New Zealand as
between hawazense and parvipedum.

Class ECHINOIDEA
Family Echinothuriidae

Hapalosoma pulchrum n. sp.
Figs. 2A-B, 3A-B

Diagnosis: A species of Hapalosoma lacking aboral primary tubercles.
Material examined: Holotype, AM J21699, 29°54.90’S, 44°.80’E, off Norfolk Island,
130-301m (NZOI stn P26).
Description: The test is flattened, more or less circular in outline, with h.d. = 67.4mm
(Fig. 2A-B).
Apical system: Measures 13mm in diameter. Ocular plates are insert, anvil-shaped,
contiguous with the more or less kite-shaped genital plates. The genital pores are large,
indicating the probability of a female. A number (8-12) of small tubercles occur on a
discrete convexity of the plates adapically to the genital or ocular pores respectively. The
madreporite is large and prominent, extending over most of the plate. Apical plates each
bear 1 or 2 tubercles (Fig. 3B).
Ambulacra: There are 55 plates in each ambulacral column. Two demi-plates, each
pierced by a pore pair, are associated with each ambulacral plate. The pore pair piercing
the ambulacral plate lies adjacent to the interambulacral plate. The pore pairs form
three columns. A large primary tubercle occurs on each second and/or third plate,
internal to the pore pairs on the oral surface as far as the ambitus of the test. A smaller
tubercle occurs on each plate between the outer demi-plate and pore-pair piercing the
ambulacral plate. A second even smaller tubercle occurs regularly under the pore-pair
piercing the ambulacral plate. The latter two tubercles form a regular double series to
the ambitus in the oral side. Three or four additional smaller tubercles occur in a
median transverse line across the plate. Above the ambitus there are no primary
tubercles but a row of 6, diminishing to 0-1 adapically, small tubercles forming a median
transverse row on the plates. Skin areas are very narrow, almost obliterated between the
plates. The ambulacral width at the ambitus is 14.2-15.6mm (Fig. 3A).
Interambulacra: There are 37 plates in each interambulacral column. The width of the
interambulacrum is about 27mm at the ambitus. A large primary tubercle occurs on
each of the first 14 plates from the edge of the peristome to the ambitus on the oral side.
These tubercles form a regular series adjacent to the ambulacra. One or two additional
tubercles, one usually close to the mid interambulacral margin of the plate, also occur,
but irregularly on these plates. Up to about twelve minute tubercles occur scattered
between the large tubercles. Above the ambitus, from about the 15th plate from the
peristome, there are no large primary tubercles. Instead, a median transverse row of
small, but equisized tubercles occur on each plate, the number diminishing from 13-14
at the ambitus to 0-1 adjacent to the apical system. Areas of skin between the plates are
minimal (Fig. 3A).

All tubercles are perforate and non-crenulate.
Peristome: The peristome measures approximately 14mm in diameter. The plates
carry flattened but club-shaped spines and pedicellariae.
Pedicellariae: These are typical of the genus though there is a tendency for the shaft of
the bifid-tipped, reduced dactylus pedicellariae to be perforated by up to three small
holes.
Colour: Test is pale green, the green being more intense on the ambulacral plates. The

PROC. LINN. SOG. N.S.W., 111 (4), 1989

264 NEW TASMAN SEA ECHINODERMS

5mm

Fig. 3. Hapalosoma pulchrum n. sp. (holotype, AMJ21699), A = plates ofambulacrum and interambulacrum, B
= apical system.

apical system is purplish. A wide purplish longitudinal band occurs in each ambulacral
and interambulacral area orally, extending just above the ambitus as a purple patch.

PROC. LINN. SOG. N.S.W., 111 (4), 1989

F. W. E. ROWE 2605

The larger oral spines are very pale greenish with 5-6 narrow purplish bands.
Remains of the white ‘hoofs’ indicate they are not prominent. The smallest secondary
spines are uniformly whitish to pale green as are the abactinal spines.

Etymology: pulcher (Lat.) = beautiful, referring to its striking colour pattern.
Remarks: This species differs from its only congeners H. pellucidum (A. Agassiz) and H.
gemmiferum Mortensen primarily by the absence of aboral primary tubercles and spines.
The pierced blade of the dactylus pedicellariae and the colour pattern may also be useful
differences separating the species. The genus Hapalosoma was last reviewed by
Mortensen (1935).

Family Pedinidae

Caenopedina alanbakeri n. sp.
Figs. 4A-B, 5A-D

Diagnosis: A species of Caenopedina with relatively short primary spines (1.37 x h.d.);
apical system 33% h.d.; peristome 25% h.d.; interambulacral plates with large primary
tubercle and prominent secondary tubercle the remaining plate surface covered with
small tubercles of 2-3 sizes; milled ring of spines 10% wider than the spine; test pale
pink, primary and secondary spines uniformly deep pink basally, lighter towards the tip.
Material examined: Holotype AM J21700, 29°24.80’S, 168°13.20’E to 20°23.70’S,
168°13.80’E, off Norfolk Island, 570-578m (NZOI stn 192).

Description: The test is circular at the ambitus, flattened aborally and orally, the sides
strongly arched (Fig. 4A-B). The h.d. = 41 mm, v.d. = 26 mm. There are 19 coronal
plates.

Apical system: This measures 13.5mm in diameter (33% h.d.) (Fig. 5A). It is dicyclic
with oculars all widely exsert. The male genital pores are small, horizontal, slit-like,
with a channel extending from the pore to the lower border of the 8th interambulacral
plate. The genital plate bearing the madreporite is not enlarged, the madreporite
occupying a triangular, central portion of the septagonal plate. There are, more or less,
two distinct groups of tubercles on each genital plate. A triangle of 10-11 tubercles
occupies the periproctal edge of the plate while a further 9-10 tubercles form a more or
less double transverse band across the plate, the remainder of the plate is bare (Fig. 5A).
The oculars bear 5-6 small tubercles of which 4-5 form a transverse line straddling the
ocular pore, the remaining tubercle occurring towards the inner edge of the plate. The
periproct is covered by small plates, the anal aperture more or less centrally placed.
Ambulacra: The ambulacra measure 6.2mm in width at the ambitus (= 35.8% of inter-
ambulacra). Ambulacral plates are trigeminate, the pore pairs being in arcs of three on
the oral surface from the peristome to the ambitus. Aborally, from about the ambitus to
the apical system the pore arcs become more vertically aligned so that the pore pairs
form a somewhat sinuous line. The middle component of each plate bears a conspicu-
ous, perforate, non-crenulate, primary tubercle whose areole extends onto the plate
components above and below it. These tubercles form a regular vertical series in each
column, decreasing in size towards the apical system. Small secondary tubercles, of two
sizes, occupy the remaining surface of the ambulacral plates (Fig. 5B).

Interambulacra: The interambulacra measure 17.3mm in width at the ambitus. The
plates each bear a large primary tubercle more or less in the middle of each plate. The
tubercles form a vertical series. The areoles are large and confluent between plates on
the oral surface. On the aboral surface, above the ambitus the areoles are separated
adapically by a simple row of small, secondary tubercles. A prominent, secondary
tubercle occurs near the mid-interradial edge of each plate, forming a second vertical
series below the ambitus but a more or less zigzag series above the ambitus. The remain-

PROC. LINN. SOC. N.S.W., 111 (4), 1989

NEW TASMAN SEA ECHINODERMS

LOE

Fig. 4. Caenopedina alanbakeri n. sp. (holotype, AMJ21700 oblique aboral view, B = lateral view (h.d. =
41mm).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

F. W. E. ROWE 267

ing surface of each plate is crowded with smaller, secondary tubercles of 2-3 different
sizes. All tubercles are perforate and non-crenulate (Fig. 5B).

imm

5mm

94um 60um Smm

Fig. 5. Caenopedina alanbakeri n. sp. (holotype, AMJ21700), A = apical system, B = plates of ambulacrum and
interambulacrum, C = milled ring of primary spine, D = spicules from tube-foot. (gg = genital groove).

Peristome: The peristome measures 10.3mm (+ 25% h.d.). It is finely plated as well as
containing the conspicuous buccal plates which bear pedicellariae.

Spines: The longest spines are 56.4mm, slender, tapering, longitudinally striated with
minute thorns. The milled ring is conspicuous but not widely produced so that it is only
10% wider than the spine (Fig. 5C). Secondary spines are similar to the primaries but
much smaller, about 1/5-1/4 the length of the primaries.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

268 NEW TASMAN SEA ECHINODERMS

Spicules: The tube feet contain spicules which are slightly curved, irregularly elongate,
perforated plates or smooth rods with marginal perforations (Fig. 5D).

Pedicellariae: These do not appear unusual for the genus, or to be distinctive, though
the ophicephalous pedicellariae are rare on the test of the holotype.

Colour: The dried test is uniformly a very pale pink (when cleaned with domestic bleach
the test is white except for a persistent pink on the plates of the apical system and
immediately adjacent coronal plates). The spines are not banded but are deeper pink
basally, becoming paler along their length, the distal 4% to % of the spine being a pale
yellowish/lime colour. The secondary spines are similarly coloured only even paler. The
poison sacks of the globiferous pedicellariae are violet/purple. The tube feet are pale
brown with the tip sienna brown.

Etymology: This distinctive species is named for Dr Alan N. Baker, National Museum
of New Zealand, who has contributed significantly to ophiuroid and echinoid taxonomy
in this geographical region.

Remarks: The genus Caenopedina A. Agassiz has been reviewed by Mortensen (1940).
More recently two species have been described from New Zealand waters; C. novaezealan-
diae Pawson (1964) and C. otagoensis McKnight (1968c). C. alanbakeri is immediately
distinguished from its geographically nearest congener C. novaezealandiae Pawson by the
colour; the size of the milled ring of the primary spines; the relative sizes of the apical
system and peristome to h.d.; the relative number of coronal plates and tuberculation of
the plates. Although the primary spines are not banded, C. otagoensis McKnight is other-
wise distinguished from C. alanbakeri on similar comparative features to those of C.
novaezealandiae. Despite the fact that the primary spines are only 1.37 x h.d., their slender
form and the test plate ornamentation immediately distinguish C. alanbaker: from either
of the short-spined C. pulchella (A. Agassiz and H. L. Clark) or C. superba H. L. Clark.
These features clearly also distinguish C. alanbakeri from all other congeners.

Class ASTEROIDEA
Family Astropectinidae

Tethyaster tangaroae n. sp.
Figs. 6A-B, 7A-B

Diagnosis: A species of Téthyaster which has stout tabulae (about twice as high as wide);
actinal plates with 1-3 prominently elongate central spinelets; adambulacral plates and
some actinal and oral plates usually bear a large, elongate bivalved pedicellaria.
Material examined: Holotype, AM J21701, 28°57.90’S, 167°45.50°E, off Norfolk
Island, 392-423m (NZOI stn P35), 2 paratypes, NZOI, 28°54.60°S, 167°44.20’E, off
Norfolk Island, 390-402m, (NZOI stn P27).
Description: The holotype measures R = 46.5mm,r = 11.3mm, br (at 2nd inferomar-
ginals) = 10.8mm; R/r = 4.1, R/br = 4.3 (Fig. 6A-B). The arms are slender, tapering
to a narrow tip which is occupied by a prominent, convex, longitudinally elongate
terminal plate. The terminal plate has a bumpy surface when cleaned of the minute
spinelets which cover it. The spinelets give the plate a shaggy, felt-like appearance. The
abactinal plates have a six-lobed base and are tabulate. They reduce in height towards
the tip of the arm. The tabulae are about twice as high as they are wide, somewhat
waisted and of two sizes, mixed, which are arranged, more or less, in longiseries on the
arms. There are thirteen series at the base of the arm, reducing to four series at the arm
tip. It is possible to detect a carinal row of spaced, larger tabulae when the tabulae are
cleaned of their spinelets. The centre of the tabulae is occupied by up to 25 short, blunt-
tipped spinelets of mixed sizes, and the periphery by up to 25 slender spinelets.

The madreporite is at about %r from the disc centre. It bears small spinelets on its

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE

Fig. 6. Tethyaster tangaroae n. sp. (holotype, AMJ21701), A = aboral view, B = oral view(R = 46.5mm).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

NEW TASMAN SEA ECHINODERMS

V),
SS, 4
AYSE
ly o
Sot

Loe
DY
isd
4
ah!

VW)
AN
U 1, Uj
NN

AY
BA,
ss
ae

y
Z¢ Yj

2mm

2
2 7
US
Y
Zz
INK
hos W Mu (NN

<— mouth

Imm

Fig. 7. Tethyaster tangaroae n. sp. (holotype, AMJ21701), A= = superomarginal (sm) and inferomarginal (im)

plates in interradial arc, B = 4th-6th adambulacral plates (ad) and adjacent actinal plates (ap), showing

spines and pedicellariae (p).

corrugated surface and is almost hidden by the spinelets of adjacent tabulae. The disc is
more or less flat, though the radial regions, at the base of the arms, are slightly convex.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 271

The papulae are single and occur between the tabulae over the whole abactinal surface,
extending to the arm tips.

The supero- and inferomarginal plates are similar in size and shape. There are 34
superomarginals and 35 inferomarginals, the distalmost 2-3 inferomarginals being very
small and extending under the terminal plate each side. The marginal plates are verti-
cally elongate, with a raised, flat-topped, rectangular median ridge which leave deep,
straight-sided fasciolar channels between successive ridges. The flat-topped surface of
the ridges are covered with low bumps, each bump bearing a short, stout spinelet. On
the superomarginal plates these spinelets become elongate towards the edge of the ridge
where together with those on the adjacent plates the spinelets span and cover the fas-
ciolar channels between. The inferomarginal plates bear a vertical series of 3,
proximally, reducing to 1 distally, large flattened, acicular spines across the ridged
surface. Additionally, smaller slender, but otherwise similar spinelets cover the remain-
ing surface with fasciolar spinelets, similar to those on the superomarginal plates, at the
periphery of the ridge (Fig. 7A).

The actinal surface is relatively small, with 4 rows of plates on each side and a
supplementary row of three plates extending along the mid-interradial line between the
oral plates and first inferomarginal plates (Fig. 6B). The first actinal row extends to
about the 10th inferomarginal plate; the second row extends to about the 6th infero-
marginal plate; the third row comprises 3 plates and the 4th is a single plate. The actinal
plates have a central convexity which bears elongate spinelets, 1-3 central ones being
more prominent, elongate, tapering spines. Several plates bear a large, elongate,
bivalved pedicellaria, replacing one or more of the central spines.

The adambulacral plates are somewhat wedge-shaped from actinal aspect, thereby
projecting, slightly, into the furrow. There are 3-5 furrow spines, the middle spine being
the longer and it is compressed. The actinal surface of the plates bear 3-4 central,
elongate spines, one or two of which are replaced by a single, large, bivalved pedicellaria
on many plates. The first two adambulacral plates each bear 2-3 such pedicellariae. The
periphery of the plates bear a number of smaller, slender spinelets (Fig. 7B).

The oral plates bear 6-7 furrow spines. The two adjacent apical spines are so closely
appressed as to appear to be fused together. Along the median crest between the adja-
cent oral plates is a row of 10-11 elongate spines, the 3rd and 4th of which are replaced by
a large bivalved pedicellaria on several of the plates.

Neither of the paratypes possess pedicellariae, otherwise they are similar in almost

all other respects to the holotype. One paratype measures R = 34.6mm,r = 9.2mm, br
= 7.6mm; R/r = 3.76, R/br = 4.6. It has 27 superomarginal and 30 inferomarginal
plates. There are 3 actinal rows and 2-3 plates form the median interradial row. The
second paratype measures R = 10.3mm,r = 4.2mm, br = 4.7mm; R/r = 2.45, R/br
= 2.2. It has 14 superomarginal and 16 inferomarginal plates. The inferomarginal
spines are hardly prominent. There are two actinal rows and 2 plates form the median
interradial row.
Remarks: T° tangaroae differs from its geographically nearest neighbour T’ aulophora
(Fisher) principally by the stouter tabulae, form of the large bivalved pedicellariae, and
spinulation of the actinal plates. 7’ tangaroae differs from the South African T’ pacez
(Mortensen) by the actinal spinulation and pedicellariae.

The discovery of a species of Téthyaster in the Tasman Sea is not an unexpected
extension in range of this widespread genus which has been revised by A. M. and A. H.
Clark (1954).

PROC. LINN. SOC. N.S.W., 111 (4), 1989

DD NEW TASMAN SEA ECHINODERMS

Fig. 8. Glyphodiscus mcknighti n. sp. (holotype, AMJ21702), A = aboral view, B = oral view(R = 23mm).

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE DIB
Family Goniasteridae

Glyphodiscus mcknighti
Figs. 8A-B, 9A-B

Diagnosis: A species of Glyphodiscus with smooth, flat abactinal plates, smooth marginal
plates and few papulae which are restricted to the radii on the disc.
Material examined: Holotype, AM J21702, 28°42.30’S, 167°56.70’E, Norfolk Island,
475-450m (NZOI stn P46).
Description: The holotype measures R = 23mm,r = 11.5 mm, br = 4.2mm (at 2nd
inferomarginals); R/r = 2.0, R/br = 5.5.

The abactinal area is pentagonal, slightly produced at the angles and sunken below
the level of the superomarginal plates. The abactinal plates are smooth, flat, without
crystal bodies. The plates are rounded-polygonal, the interradial plates being slightly
larger than the radial plates. A row of narrow, transversely rectangular plates occurs
adjacent to the superomarginals. A small, triangular madreporite occurs 4%r from the
centre of the disc (Fig. 8A).

The supero- and inferomarginal plates are similar in shape, size and number. They
are block-like, smooth, longer than wide with a rounded dorsal-lateral or actinal-lateral
edge respectively. There are 6 of each on each side of the disc and arm, the first 2 supero-
and inferomarginals of adjacent radii respectively delimiting the pentagonal abactinal
and actinal disc surfaces. The remaining 4 supero- and inferomarginals on each side of
the arms unite across the arm along the median line. The terminal plate is small, with a
convexity at its tip on either side of the actinal channel which houses the terminal tube
foot. This channel is guarded by 3-4 minute granules.

The actinal plates are flat, smooth, rather transversely diamond-shaped. They
form a regular pavement arrangement (Fig. 8B).

The abactinal, marginal and actinal plates are each surrounded by a single row of
minute granules so that a double row of granules occurs between adjacent plates.

Papulae are restricted in each radius abactinally and are delimited as 5 rounded,
convex areas each comprising about 10 plates.

Usually 1, occasionally 2, small spatulate pedicellariae occur at the edge of a
number of actinal plates, also at the actinal edge of several of the inferomarginal plates
(Fig. 9A). Pedicellariae are not present on the superomarginal or abactinal plates of this
specimen.

The adambulacral plates bear five laterally compressed furrow spines (Fig. 9B),
behind which stands a row of 3 enlarged (subambulacral) granules. The remaining
actinal surface of the plate bears 2-3 rows each of 3-4 smaller wedge-shaped granules
which merge in size with those surrounding the actinal plates (Fig. 9A).

The oral plates bear 7-8 furrow spines, 4-5 enlarged (suboral) granules and a tri-
angular group of 8-10 wedge-shaped granules on the remaining actinal surface of the
plate.

Etymology: Named for Mr D. McKnight of NZOI who has described much of the
Tasman echinoderm fauna.

Remarks: I have placed this new species in the herein elevated subgenus Glyphodiscus
Fisher on the grounds that the species characters are consistent with those outlined for
the genus by Fisher (1917, 1919). I believe the smooth plates which lack glassy bumps
(crystalline bodies), the complete ring of peripheral granules around all plates and the
form of the adambulacral armature are quite reasonably sufficient to distinguish
members of this genus from those in either Jconaster Sladen, Lithosoma Fisher or Astro-
ceramus Fisher.

G. mcknighti is clearly distinguished from its only congener G. perzerctus Fisher which

PROG. LINN. SOC. N.S.W., 111 (4), 1989

274 NEW TASMAN SEA ECHINODERMS

has roughened superomarginal plates, tumid peripheral abactinal disc plates and
papulae distributed over the disc. In respect of size and shape, the two species are very
closely similar.

2mm

1mm

— mouth 4th ad

Fig. 9. Glyphodiscus mcknighti n. sp. (holotype, AMJ21702), A = spine and granule arrangement on
adambulacral (ad) and adjacent actinal (ap) plates with pedicellariae, B = furrow spines (fp).
Family Brisingidae

Novodinia helenae

Figs. 10A-B, 11A-C

Diagnosis: A species of Novodinia with 12 arms, disc with papulae in groups of up to 10,

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 275

single madreporite, membranous, unskeletonised genital region of arm, and single
adambulacral spine.

Material examined: AM J21703, 29°20.20’S, 168°10.79’E, off Norfolk Island, 308m
(NZOI, stn 194).

Description: The holotype has 12 rays; dd = 14mm, R = 75mm, br = 3.8mm (at
base), 5.5-5.9mm (at widest part of genital expansions 12mm from base of ray), 3.0mm
(at %R); R/r = 10.7 (Fig. 10A-B).

The disc is circular, 3.7mm high, flattened abactinally. It is covered by overlapping
scale-like, and convex abactinal plates which form an open reticulum in which groups of
up to 10 (occasionally single) papulae occur. Sharply pointed spines (up to 1.2mm),
wreathed almost to their tips with crossed pedicellariae (Fig. 11C) occur on a number of
the convex plates. These spines may stand singly, in pairs or triplets. In the latter two
cases the spines are united by a web across which the pedicellariae span.

The genital region, at the base of the arm, is covered by a thin, unskeletonised
membrane. This region is crossed by 4-5 complete costae. The costae comprise the
marginal plate each side of the arm, linked more or less regularly by 7-10 rod-shaped
abactinal plates. Costae 1 and 2 may be irregularly linked by a few abactinal plates
extending mid-dorsally between them. Most of the plates of the costae, except for the
marginal plates of the first two costae, bear a single, sharply pointed, slender spine.
Most of these spines are encased in a gland-like sheath of pedicellariae. Beyond the 4th-
5th costae the spine bearing marginal plates and abactinal plates form incomplete
costae, that is to say a marginal and 4-5 small abactinal plates project dorsalwards along
the side of the arm on each side but do not link over the abactinal surface. The link is
made instead by a wide band of crossed pedicellariae. The abactinal surface of the arm
is then covered by a very thin membrane for the rest of the length of the arm. Abactin-
ally, between each of the incomplete costae a rounded patch of pedicellariae also occurs
along the arms. The incomplete costae occur opposite every 4th-5th adambulacral plate
along the arm (Fig. 11A).

The adambulacral plates are block-like, wider than long and each bears a large,
cylindrical spine (2.5mm) which at least for the length of the genital area are flared into
a bi-quadrifid tip. Beyond the genital area these spines are slender and pointed. These
spines bear numerous pedicellariae, but these are confined to the outer surface and do
not ensheath the spines. The first adambulacral plate of each arm is fused across the
interradial line to its adjacent neighbour. However the first two adambulacral plates on
each side of each arm are united by a syzygy. Thereafter the plates are united by muscle
blocks, the interstices between each successive pair of adambulacral plates being half the
length of the plates themselves.

The actinosome is 8mm in diameter and the mouth is 3.6mm wide. The peristome
is thin but translucent. The oral plates are as usual for the genus, with lateral processes
meeting mid-radially. In well-developed oral angles each plate bears 5 spines in a fan-
shaped, marginal arrangement. The innermost spine is usually very small, rarely is it
long and bearing pedicellariae. The second spine is always long and pointed and bears
pedicellariae. The remaining 3 spines are small, non-pedicellariae bearing and decrease
in size towards the furrow. Where arms are being regenerated, the mouth angle plates
bear fewer spines (3-4) of which the elongate pedicellariae bearing spine is in the apical
position. A small additional spine can be seen to develop in the interradial side of this
spine in angles with more advanced regeneration. There are no spines on the actinal
surface of the oral plates. The furrow margin of each oral plate is excavate to accommo-
date the first pair of tube feet (Fig. 11B).

The madreporite (1.2mm diameter) is small, subtuberculate, coarsely furrowed.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

276 NEW TASMAN SEA ECHINODERMS

Fig. 10. Novodinia helenae n. sp. (holotype, AMJ 21703), A = aboral view, B = oral view (dd = 14mm, R = 75
mm).

PROC. LINN. SOG. N.S.W., 111 (4), 1989

F. W. E. ROWE BT

7mm
120pm

ad ads

Fig. 11. Novodinia helenae n. sp. (holotype, AMJ21703), A = costae (c) in genital region of arm, B = oral plates
(op) and arm bases with spine arrangements, C = value of crossed pedicellaria (ad = adambulacral plate;
ads = adambulacral spine; tf = tube-feet).

Etymology: Named for Dr Helen E. S. Rotman (nee Clark) who has contributed to our
knowledge of Tasman and Antarctic asteroids.

Remarks: The new species clearly belongs in the genus Novodinia Dartnall et al., as most
recently redefined by Downey (1986). Only two species of Novodinia are recorded from
the Tasman/New Zealand region. N. australis (H. L. Clark), from southeastern Aus-
tralian waters and N. novaezealandiae (H. E. S. Clark), from off the Chatham Islands, east
of New Zealand. N. helenae is immediately distinguished from each of these species by a
number of characters: firstly, low arm number (12) as opposed to 14-16 (australis) or 18
(novaezealandiae); secondly, abactinal skeletal arrangement of the disc with groups of
papulae (in each of australis and novaezealandiae the skeletal network is close and papulae
occur singly); thirdly, the genital area of the arm is membraneous in N. helenae but plated
in the other two species. The number of madreporites (1 in Aelenae but 4 in australis) and
the number of adambulacral spines (1 in Aelenae but 3-4 spines in novaezealandiae) are also
features of distinction. Arm number and ornamentation of disc and genital regions
readily distinguish N. helenae from its other congeners.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

278 NEW TASMAN SEA ECHINODERMS

Class CRINOIDEA
Family Antedonidae

Nanometra duala n. sp.
Fig. 12A-E

Nanometra johnstont. McKnight, 1977: 136 (non N. johnston: John)
Diagnosis: A species of Nanometra with a pair of wing-like extensions to the segments of
P, and subsequent pinnules.
Material examined: Holotype, AM J21704 and 2 paratypes, NZOI, 29°20.20’S,
168°10.79’E, off Norfolk Island, 308m (NZOI, stn 194).
Description: The holotype has 10 arms broken at about 20br and measuring about
10mm in length. The estimated length of the arms is not more than 20mm (Fig. 12A).
The cirri are XXIV-X XVI, 19-22. The first 2 segments are twice as broad as they are
long, the third segment is 1.5 x as long as it is broad, 4th-5th segments twice as long as
broad, 6th segment 1.75 x as long as broad, 7th segment 1.25 x as long as broad. Distally,
the segments are as long as they are broad. The segments do not bear a dorsal spine but
the distal end is expanded (Fig. 12A-B).

The centrodorsal is conical, with a rugose apex. The cirri are arranged in vertical
rows of 2 or 3 (Fig. 12B).

The radials are narrow, almost hidden, with a tubercle at each of the exposed
corners of the plate.

The 1Br, is twice as broad as it is long, with the proximal and distal edges more or
less straight but with the distal edge everted. There is a rounded or chisel-shaped
tubercle arising on each side of the ossicle and one or two spinulose tubercles below each
of these on the dorsal-lateral surface of the ossicle. The axillary (1Br,) is triangular,
slightly broader than long (1.25:1). The proximal border is slightly convex, the distal
border everted and spinulose. There are one or two spinulose tubercles occurring on the
lateral surfaces of the ossicle (Fig. 12B-C).

Br, is quadrate, about 2-2.5 times as wide as long. Up to 4 spinulose tubercles may
occur on the lateral surface of the ossicles. Remaining brachials are more or less
elongate, wedge-shaped, with the distal edge everted and spinulose (Fig. 12C-D).
Syzygies occur at 3+4 and usually 9+10, but occasionally the second syzygy occurs at
8+9 or 11+12 or 13+14. The third syzygy occurs at 14+15 or 15+16. P, is the longest
pinnule and the stoutest. It is 3mm long, comprising 11 segments. The first segment is
broader than long, the second is quadrate, the remaining segments are up to 2 times as
long as broad. P,, is about 1.5mm long and comprises 6 segments. P, is smaller and more
slender than P,, is about 2mm long and comprises 9 segments. The segments of these
pinnules are strongly everted and spinulose on their distal edge. P, is the first gonadal
pinnule. On P, and subsequent pinnules (for the length of the broken arms) the lateral
edges of the 2nd to 4-5th segments are expanded into thin, wing-like processes, giving
the pinnules a very characteristic form (Fig. 12D).

One paratype is very similar indeed to the holotype in being a relatively intact, if
broken armed, specimen. The second paratype comprises the calyx of a specimen with
many broken arm pieces. The form of the segments of the pinnules is, however,
unmistakable in uniting the specimens within a single species.

Etymology: duo = two; ala = wing (Lat.) referring to the 2-winged appearance of the
pinnular segments.

Remarks: The very distinctive form of the pinnule segments immediately distinguishes
this species from its congeners N. johnstoni John (from S.E. Australia); N. clymene A. H.
Clark (from the East Indian region) and N. bowers: (A. H. Clark) (from southwestern
Japan), which have been reviewed by A. H. Clark and A. M. Clark (1967). McKnight’s

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 279

(1977) record of N. johnston: from the same locality (NZOI stn 194) refers instead, I
believe, to the new species N. duala.

Smm

2.5mm

2mm

Fig. 12. Nanometra duala n. sp. (holotype, AMJ21704), A = lateral representation of holotype, B = cirri,
centrodorsal and 1Br, C = 1Br and arm bases, D = P3 with cross-section of pinnule segment showing wing-
like extensions.

ACKNOWLEDGEMENTS

The author wishes to express his gratitude to the following people for allowing him
unlimited access to their collections and for the loan of material: Messrs D. McKnight
and W. L. Main (New Zealand Oceanographic Institute, Wellington); Drs A. N. Baker
and H. E. S. Rotman (National Museum of New Zealand, Wellington); Sue Boyd
(Museum of Victoria, Australia). Ms P. Berents and Ms Lynne Albertson (Australian
Museum) are thanked for preparing figures and Ms Berents for reading and comment-
ing on the manuscript. Mr. A. Farr (Australian Museum) for photographing the
specimens. Ms Jan Howarth is thanked for typing the manuscript. This project has been

PROC. LINN. SOC. N.S.W., 111 (4), 1989

280 NEW TASMAN SEA ECHINODERMS

funded by Marine Sciences and Technologies Grants Scheme (MST 84/2092) which is
duly acknowledged.

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PROC. LINN. SOC. N.S.W., 111 (4), 1989

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PROC. LINN. SOC. N.SW., 111 (4), 1989

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PROC. LINN. SOC. N.S.W., 111 (4), 1989

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PROC. LINN. SOC. N.S.W., 111 (4), 1989

A Review of the Family Caymanostellidae
(Echinodermata: Asteroidea) with the
Description of a New Species of
Caymanostella Belyaev and a New Genus

FRANCIS W. E. ROWE

Rowe, F. W. E. A review of the family Caymanostellidae (Echinodermata: Asteroidea)
with the description of a new species of Caymanostella Belyaev and a new genus.

Proc. Linn. Soc. N.S.W. 111 (4), 1989: 293-307.

The history of the deep-sea asteroid family Caymanostellidae is reviewed. A
recently proposed close relationship between Caymanostella and the concentricycloid
genus Xyloplax is discussed and refuted. A new species in the genus Caymanostella is
described from the Tasman Sea and a new genus, Belyaevostella, is described to
accommodate the Indonesian species C. hispida Aziz and Jangoux. New observations
for caymanostellid asteroids are the occurrence of papulac in the new genus, varied
position of the gonopore associated with the proximal-most superomarginals and
internal structure of some abactinal plates.

Francis W. E. Rowe, Division of Invertebrate Zoology, Australian Museum, PO. Box A285, Sydney
South, Australia 2000; manuscript recewed 7 December 1988, accepted for publication 19 July
1989.

INTRODUCTION

The family Caymanostellidae was described by Belyaev (1974) to accommodate the
then unique species Caymanostella spinimarginata Belyaevy, collected from depths of 6740-
6780m in the Cayman ‘Trench and living on sunken wood. Belyaev included the family
in the asterozoan order Phanerozonia because of the prominence of the marginal plates.
He was uncertain whether the distinctive features including the shape of the oral plates,
lack of ventro-lateral (actinal) plates, and the actinal spinulation represent primitive
(plesiomorphic) features or derived (apomorphic) characters related to the specific
habitat of the sea-stars. Consequently, he was also uncertain of relationships of the
Caymanostellidae with other phanerozone families.

Later (1977) Belyaey and Litvinova described a second species, C. admiranda, which
had been collected in 5220m from the northern part of the Coral Sea, West Pacific. They
made no further comment on the relationships of the family.

Aziz and Jangoux (1984) have described a third species C. hispida, from 2350m in
the Strait of Macassar (Indonesia), West Pacific. They suggest that because of the form
of the abactinal and marginal skeleton, the family has closer affinities with members of
the order Spinulosida, and in particular the family Asterinidae.

Smith (in Smith and Tranter, 1985) described a fossil sea-star, Protremaster unisertalts,
from Sinemurian (Lower Jurassic) deposits on Alexander Island, Antarctica. He con-
sidered the species a tremasterin asterinid, but compared it, also, with Caymanostella.
Rowe e al. (1988) and Rowe (1988), however, consider Protremaster to be an early
caymanostellid asteroid, thus concluding that the family Caymanostellidae is of
relatively ancient lineage and that early members possessed at least one row of actinal-
lateral plates.

Blake (1987), in a major reclassification of post-paleaozoic asteroids, refers the
Caymanostellidae to the order Velatida, which he re-established to include also the
Korethrasteridae, Solasteridae, Myxasteridae and Pterasteridae. The Velatida are

PROC. LINN. SOC. N.S.W., 111 (4), 1989

294 A CAYMANOSTELLID ASTEROID REVIEW

included with the monofamilial Spinulosida within the superorder Spinulosacea. Blake
(1987) contends that the Caymanostellidae are aberrant velatidans and that characters
shared between Caymanostella and members of the Valvatida are ‘. . . best explained by
convergence.’ Unlike Aziz and Jangoux (1984), Blake (1987) does not draw a direct
comparison with the Asterinidae, which latter family he had earlier (1981) transferred
from the Spinulosida, (e.g. Spencer and Wright, 1966) to the Valvatida. Rowe et al.
(1988) maintain the Caymanostellidae, Korethrasteridae and Asterinidae should be
retained within the same order, the Valvatida. Rowe et al. (1988), in discussion of the
origin of the Class Concentricycloidea, suggest that the caymanostellids and korethras-
terids may warrant a supra-familial taxon of their own.

Smith (1988) classifies the Caymanostellidae as a sister group to Xyloplax
medustformis Baker, Rowe and Clark (1986) within the order Velatida (sensu Blake, 1987).
He argues against the recognition of the Class Concentricycloidea erected by Baker et al.
(1986) for Xyloplax.

In this paper, and following examination of over 250 specimens of caymanostellids,
the family Caymanostellidae is comprehensively re-diagnosed and its relationships,
including that with Xyloplax are discussed. A new species of Caymanostella is described
and a new genus described for C. hispida Aziz and Jangoux. Two paratypes of C.
spinimarginata Belyaev, one paratype of C. admiranda Belyaev and Litvinova and the
holotype and paratype of C. hispida Aziz and Jangoux have been re-examined. In the
text, Institution abbreviations are AM = Australian Museum; BM(NH) = British
Museum of Natural History, London; NMNZ = National Museum of New Zealand,
MNNH = National Museum of Natural History, Paris.

SYSTEMATIC ACCOUNT
Family CAYMANOSTELLIDAE Belyaev

Caymanostellidae Belyaev, 1974: 1502

Diagnosis (emended): Body pentagonal to circular in outline; aborally slightly convex,
orally flat; body covered with thin or very thick epidermis; madreporite relatively
simple; disc plate arrangement distinct or not, from arms, aborally; disc plates variously
imbricated include centrodorsal, primary radial and interradial plates, a pair of distinc-
tive distal-lateral disc plates (dldp) in each interradius and several intercalary plates;
each dldp has an internal, horn-like process which abuts with its opposite number in the
mid-interradial line, forming, with the oral plates, internal buttressing points between
which a [shaped odontophore is braced (Figs. 1, 5F); aboral arm plates imbricate
proximally, comprising a carinal and one or more dorsal-lateral rows each side; aboral
plates hexagonal, to fan-shaped, thin delicate and scale-like or thicker lenticulate and
robust; superomarginals not conspicuous in fossil taxon but more or less rectangular
conspicuous in Recent taxa, entirely abactinal, first pair usually markedly larger than
succeeding ones, supermarginal plates either similar in size or smaller than infero-
marginal plates; inferomarginals rectangular, delimiting margin of body at oral/aboral
angle; terminal plates each with central perforation; actinal-lateral plates absent in
Recent forms, present in known fossil taxon; adambulacral plates bar-like, extending
between ambulacral and inferomarginal plates in Recent taxa; first adambulacral plate
very short, second and third longest; furrow margin straight, furrow spines few (1-3);
ambulacral grooves narrow or broad/petaloid; two rows of suckered tube-feet; 10
gonads, 1| pair to each interradius, visible from oral surface through thin, membranous,
triangular ‘window, actinal membrane with or without perforated plate spicules (Fig.
7); 10 gonopores one pair to each interradius, associated with proximalmost supero-
marginal plate of each plate series, as a marginal notch or piercing the plate, or

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 295

0.5mm

Fig. I. Internal juxtaposition of horns of distal lateral disc plates (dldp), odontophore (0) and oral plate (op),
viewed from oral surface.

positioned adjacent to first superomarginal plate (Fig. 2) (position of gonopores not
determined in Fossil taxon); sexually dimorphic, male gonopores minute compared with
size of female gonopores; abactinal plates bear granuliform or long, slender spinelets;
adambulacral plates bear spike-like spinelets, inferomarginal plates bear 1-3 short,
clavate or slender spines on their outer edge, forming an ambital fringe to the asteroid;
no pedicellariae; papulae present or absent; habitat of Recent taxa sunken, waterlogged
wood; Lower Jurassic — Recent.

Type genus: Caymanostella Belyaev, 1974 (own designation).

Other general included: Protremaster Smith, 1985; Belyaevostella gen. nov.

Remarks: The Caymanostellidae share with the Korethrasteridae Danielssen and
Koren, 1884, the form of the adambulacral plates, the occurrence of an actinal
membrane, though this latter is better developed in caymanostellids, and the absence of
actinal-lateral plates in Recent taxa. Lack of fossil material of korethrasterids prevents
comparison of early actinal morphology between the two families. Both families have
taxa which possess abactinal papulae and taxa which do not. They differ, however, in
shape, korethrasterids are stellate whereas caymanostellids pentagonal; abactinal
plating, (3 of 4 genera of korethrasterids — Peribolaster Sladen, 1889, Remaster Perrier,
1894 and Anareaster, Fell and Clark, 1959) have cruciform plates; abactinal spination,
korethrasterids have paxilliform plates whereas caymanostellids have an even covering
of granuliform spinelets or spaced, elongate spinelets; and the more prominent develop-
ment of marginal plates in caymanostellids.

Unless the form of the adambulacral plates and occurrence of an actinal membrane
in both families can be shown to be the result of convergence, then they may be con-
sidered to be related, commonly derived and included within the same asteroid order.
As far as the caymanostellids are concerned, the form of the Fossil Protremaster indicates
Recent forms have lost actinal plates, though the adambulacral structure was already
evolved in the Jurassic genus. There is no evidence to suggest that the bar-like adam-
bulacral plates resulted from fusion with actinal-lateral plates. It can only be speculated
that the extension of the adambulacral plates and loss of actinal-lateral plates is a
response to specialized habitat/environmental pressures.

PROG. LINN. SOG. N.S.W., 111 (4), 1989

296 A CAYMANOSTELLID ASTEROID REVIEW

A B C

Fig 2. Position of gonopores in Caymanostella (A = marginal, B = piercing) and Belyaevostella (GC = inter-
proximal-most superomarginals).

Smith (1985) clearly saw in Protremaster close affinities with asterinid taxa rather
than with Caymanostella. Paradoxically, therefore, there is a measure of agreement
between Smith (1985) and Rowe ef a/. (1988) regarding the relationship of caymano-
stellids and asterinids when Smith concludes Caymanostella may be a specialized offshoot
from tremasterins. I would support this proposition in referring to the possible
derivation of caymanostellids from asterinids since I am unconvinced of the subfamilial
relationship of tremasterins within the family Asterinidae (Rowe and Berents in prep.)
and Smith’s genus is not a tremasterin in any case. This view places the families
Caymanostellidae and Korethrasteridae with the Asterinidae in the order Valvatida
under Blake’s recent reclassification. Whether a separate supra-familial taxon is
warranted (Rowe, 1988) for this group of families, remains to be determined.

However, I would contend that Smith’s (1988) comparison of Caymanostella and
Xyloplax contains errors of interpretation due to his reliance on inadequately detailed
published data and lack of examination of relevant specimens. Full descriptions of
Xyloplax spp., including sperm development and morphology, are now available (Rowe et
al., 1988; Rowe, 1988; Healy et al., 1988). The Caymanostellidae are detailed herein. It
can now be shown that a number of characters chosen by Smith (1988), to show a close
relationship between Caymanostella and Xyloplax, are incorrect (see Table 1). I would also
point out that caymanostellids reach a diameter of at least 27mm (R = 13.6mm in
Belyaevostella hispida), three times the 9mm recorded by Smith (1988: 20).

There can be, in my view, no support for a progenetic origin for Xyloplax (Smith
1988: 20) based on the early form of the asteroid water vascular system. Smith contends
that in metamorphosing asteroids, before arm and ambulacral skeletal development
‘. .. the first few tube feet in effect form a peripheral circle’ In my view, this would be
due to the juxtaposition of a small, primary (terminal) tube-foot adjacent to which, and
on either side, are one or two larger ambulacral tube feet developing along a primordial
radial canal. There is no evidence of an incipient double-ring arrangement as evolved in

Xyloplax. Such an arrangement, if it occurred, would have profound implications in the
development of the radial structure of the ambulacral water vascular system in adult
echinoderms. There is no embryological evidence that such is the case in asteroids or
any other echinoderms.

Contrary to Smith’s (1988) reliance on the superiority of his cladistic approach to
the relationship between Caymanostella and Xyloplax, 1 do not believe that his use of
cladistics has increased the objectivity of approach to this particular problem. In my
view the relationship between Caymanostella and Xyloplax remains as remote as that
described by Baker et al., 1986; Rowe et al., 1988; Rowe, 1988 and Healy et al., 1988.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 297

TABLE 1

Comments on characters chosen by Smith (1988) in cladistic analysis

Character chosen by
Smith (1988)

Calcinal ring

2. Superomarginals
3. Hyaline spines
4. Ambulacrals
5. Oral plates
6./14. Adambulacral
arrangement
7. Actinals
8. ‘Terminal plates
9. Inferomarginals
10. Peristome/oral plates
11. Madreporite
12. Abactinal plates/
papulae
13. ‘Tube-feet
14.
15. Ambulacral plating

Comments

Inappropriate for use above generic if not species level.

Palaeontological evidence does not support Smith’s use of this character in such a
narrow sense.

Highly derived; presence or absence of these specialized spines is too widespread
in the Asteroidea for use to compare the Concentricycloidea. In any case they
are absent in both caymanostellids and xyloplacids.

Interpretation is critical. Rowe et al., (1988) and Rowe (1988) interpret
ambulacrals of Xloplax as having fused with the adambulacral/oral plate to form
the expanded oral frame.

Interpretation is critical. The oral plates are not unusual in the asteroid
Caymanostella, but substantially altered in Xyloplax, forming, with fusion to
ambulacral plates a uniquely widened oral frame (Rowe et al., (1988); Rowe
(1988) ).

In caymanostellids the adambulacrals, except the first one, have extended in
width across the actinal surface. The first adambulacral is NOT reduced to

rudimentary state as claimed by Smith (1988: 20). The adambulacrals are NOT
lost from Xyloplax but have changed roles to act as ‘ambulacral’ plates. Articu-
lation between the adambulacrals and the ambulacral/oral frame ossicles is
substantially altered (see Rowe et al., 1988).

These are present in caymanostellids, even if only in the fossil taxon, Protremaster,

described by Smith (1985).

Perforated terminal plates occur across a diverse range of taxa (Rowe, 1985)
including Caymanostellids, but not Xyloplax.

These are differentiated in each of the groups included in Smith’s (1988)
cladogram.

The oral frame of Xyloplax comprises a ring of ossicles formed by fusion of
adambulacral oral/ambulacral plates.

Caymanostellids have a small but typical madreporite.

One caymanostellid (described herein) DOES possess papulae; the type genus of
the korethrasterids, Korethraster, does NOT possess papulac; plates are imbricate,
not tessellate; caymanostellids bear granuliform and spiniform abactinal
armament on plates.

Caymanostellids possess suckered tube-feet NOT pointed tube-feet. This was a
mistake made by Belyaev (1974) and perpetuated by Smith (1988) clearly
without reference to any specimens. The pointed tube-feet of Xyloplax are
explained by Rowe-(1988) to be probably functionally associated with living in a
soft substrate.

See 6.

A critical difference in interpretation by Rowe ef al. (1988) and Rowe (1988) who
consider the perradial plates between which the tube-feet protrude to be
modified adambulacrals.

Genus Caymanostella Belyaev

Caymanostella Belyaev, 1974: 1502

Diagnosis (emended): Caymanostellid sea-stars with hexagonal to fan-shaped, lenti-
culate abactinal plates; thin epidermis not obscuring skeletal plates (in adults); granuli-
form abactinal armament; ambital spinelets club-shaped; gonopores form a notch in
aboral edge of plate (Fig. 2A) or pierce the first superomarginal plates (Fig. 2B); spicules
absent from actinal membrane; papulae absent, distributed in the Caribbean and West
Pacific.
Type species: C. spinimarginata Belyaev, 1974, by own designation.

Other species included: C. admiranda Belyaev and Litvinova, 1977; C. phorcynis sp. nov.

PROC. LINN. SOC. N.S.W., 111 (4), 1989

298 A CAYMANOSTELLID ASTEROID REVIEW

Remarks: Caymanostella is distinguished from Belyaevostella gen. nov., the latter genus
possesses delicate, scale-like abactinal plates embedded in a thick epidermis (in adults),
spiniform abactinal and marginal armament, abactinal papulae and spicules in the
actinal membrane. The gonopores of Belyaevostella occur adjacent to but do not pierce or
notch the superomarginal plates.

Caymanostella spinimarginata Belyaev
Figs 2A, 3A-B

Caymanostella spinimarginata Belyaev, 1974: 1504, figs 1-2.

Material examined: 2 specimens (paratypes, BM(NH) 1987.3.26.1 (R = 4.8mm) and
1987.3.26.2 (R = 4.3mm) ), 19°38.5’N, 76°37.5"W, 6740-6780m (Oriente depression,
eastern part of Cayman Trench, Stn 1267), R/V ‘Akademik Kurchatov’, 25.3.73.
Diagnosis: A species of Caymanostella in which the abactinal plates imbricate in a similar
direction, only the primary interradial plates (including the madreporite) are distin-
guishable on the disc; there are at least two alternating, dorsal-lateral rows of plates
either side of the carinal row along each of the arms; the supero- and inferomarginal
plates are of similar size to each other; the gonopore is positioned in a notch on the upper
margin of the proximal-most superomarginal plate in each series; the oral plates each
with 3 oral, and a single suboral, coarsely thorned spines; adambulacral plates with a
single furrow and up 10 coarsely thorned spines, these latter mostly in two alternating
rows along the actinal surface of the plate; outer edge of inferomarginals each with 2
elongate, club-shaped spines, actinal membrane triangular to heart-shaped; abactinal
granulation homogenous, blunt, club-shaped; maximum known size for species R =
4.8mm; distribution, Cayman Trench.

Remarks: The marginal position of the gonopore on the proximal-most superomarginal
plates immediately separates this species from C. admiranda Belyaev and Litvinova and
C. phorcynis sp. nov. Differences in abactinal plate arrangement also-separate the species.

Caymanostella admiranda Belyaev and Litvinova
Fig. 4A-B
Caymanostella admiranda Belyaev and Litvinoya, 1977: 1983, figs 1 and 2.

Material examined: 1 specimen (paratype, BM(NH) No. 1987.3.26.3 R = 3.4mm)
VE SMM AS ESET, (CorilSee)), D2 A0rn, RAY, Dron IWilemelelea (Sinn, W254), WIN2.7/5s 2
specimens (Amsterdam Museum, R = 4mm; R = 2.1mm) 10°39’S, 123°40’E (off
Timor) 520m, (soft grey mud with brown upper layer), ‘Siboga Expedition, (Stn. 297).

Diagnosis: A species of Caymanostella in which the abactinal plates imbricate and are ar-
ranged in two distinct fields; on the disc the centrodorsal plate is surrounded by an inner
ring of 7 plates, 5 large trapezoidal radial plates and, in each of 2 of the interradu, a
small squarish plate, there is an outer ring of 10 plates, 5 large, hex-heptagonal inter-
radial plates and 5 smaller hexagonal, rather shield-shaped radial plates; between the
primary interradial plate and radial plate on the outer ring of the disc plates, and the
proximal carinal plate and dorsal-lateral plate is an obliquely directed, oblong distal-
lateral disc plate; along the arms are 3 heptagonal-fan-shaped carinal plates, with 2-3
dorsal-lateral plates alternating either side; the terminal plate is radially longer than
wide, with a relatively small perforation towards its distal edge; there are 6-7, more or
less equal-sized superomarginal and inferomarginal plates, with the exception that the
proximal-most superomarginal in each series is about twice as broad as its immediate
neighbour, a small, wedge-shaped plate separates the upper half of the proximal-most
two superomarginals across the mid-interradial line: a small gonopore (male) pierces
the centre of each of the first superomarginal plates; the oral plates each bear 2 oral and

PROC. LINN. SOG. N.S.W., 111 (4), 1989

FE. W. E. ROWE 299

Fig. 3. Caymanostella spinimarginata Belyaev (paratype, BM(NH) 1987.3.26.1). A = aboral view, B = oral view
(R = 4.8mm).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

300 A CAYMANOSTELLID ASTEROID REVIEW

Fig. 4. Caymanostella admiranda Belyaev and Litvinova (paratype, BM(NH) 1987.3.26.3). A = aboral view, B
= oral view(R = 3.4mm).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

F. W. E. ROWE 301

one suboral, rather coarsely thorned spines; adambulacral plates bear a single furrow
spine and only one or two coarsely thorned spines on the actinal surface of the plate; the
outer edge of the inferomarginal plates bear 2 marginal spines; the actinal membrane is
heart-shaped; furrows are petaloid with up to 8 pairs of tube-feet; abactinal granulation
even, over abactinal surface, granules low, rounded, 2 sizes more or less apparent;
maximum known size, R = 4.90mm,; distribution Coral and Timor Seas.

Remarks: Belyaev and Litvinova (1977) neither described nor adequately figured the
abactinal plate arrangement for their species. The diagnosis, therefore, includes infor-
mation based on one of the two paratype specimens which has been donated to the
British Museum (Natural History) in London by Dr. G. Belyaev. Both of the ‘Siboga’
specumens (R = 4mm, r = 3.3mm, R/r = 1.2mm; R = 2.1mm, r = 1.8mm, R/r =
1.16) are in very poor condition, broken, and have apparently dried out at some time
prior to this examination. The specimens are currently stored in alcohol, but dissoci-
ation of some plates indicate probable original storage in formaldehyde solution.
Because of the fragility of the specimens, the abactinal granules have not been removed
to examine the plate arrangement. However, where granules have already been lost, it is
evident that the hex-heptagonal shape of the plates and abactinal plate arrangement is
similar to that described for C. admiranda. This second record for C. admiranda therefore
requires confirmation when better preserved material becomes available from the
Timor Sea, particularly considering the shallow depth from which the specimens were
collected.

The shape and arrangement of the abactinal plates clearly identifies C. admiranda
from its congener C. phorcynis sp. nov., with which it shares the feature of the gonopores
piercing the superomarginal plates. Currently, the geographical range of C. admiranda is
considered to extend between the Coral and Timor Seas in depths of 5220m and 520m
respectively.

Caymanostella phorcynis sp. nov.
Figs. 2B, 5A-D

Material examined: 3 specimens (holotype, AM J18911, 2 paratypes, (1 dissected for
SEM)VAM, J17880) 34°54°S, 151°14 Eto 34°507S, 151°15’E, 993-1030m (off Shoal-
haven Bight, New South Wales, Australia). N.S.W. State Fisheries R/V ‘Kapala’,
26.10.83; 3 specimens (paratypes, AM J18910 (1), AM J17830 (2) ) 35°31’S, 150°50’E to
35°28’S, 150°53’E, 938-994m (off Shoalhaven Bight, New South Wales, Australia),
N.S.W. State Fisheries R/V ‘Kapala, 8.9.83; 2 specimens (paratypes, AM J16365)
33°40’S, 151°56’E to 33°37’'S, 151°56’E, 736m (off Broken Bay, New South Wales,
Australia), N.S.W. State Fisheries R/V ‘Kapala’ 6.12.77; 9 specimens (paratypes NUNZ
Bet c(o) ee ON en OA (2) PANN 18913 (2) ie Sie 23a) Sl 230 De Beton ye 2327 (9;
177°36.6’E, 1075-1100m (off White Is. north island of New Zealand), USSR f/v
‘Kalinovo’”, on/in large waterlogged log of Corzaria arboraea. 3 specimens (paratypes,
NII AAD Ae 4224 2223.) 1794s 247 2 °W, 41S Ona BONE: of
Chatham Islands), F-V. ‘Otago Buccaneer’, 22.7.84, on wood; 230+ specimens (NMNZ
Sop ele Sualii 395) 1 to) 3 2 3efa Syl 3 6.028,” 1075-11 00m (ei VWihite Ts;
north island of New Zealand), USSR f/v ‘Kalinovo, on/in large waterlogged log of
Coriaria arborea; 7 specimens (Amsterdam Museum) 3°27.1°N, 125°18.7’E, 2053m, fond
dur, traces de sable fin, foncé et dur, ‘Siboga’ Expedition (Stn. 126).

Diagnosis: A species of Caymanostella with fan-shaped plates; gonopores piercing the
proximal-most superomarginal plates; 8-15 supero- and inferomarginal plates similar in
size (except larger proximal-most superomarginals); ambulacral furrows narrow;

PROC. LINN. SOG. N.S.W., 111 (4), 1989

302 A CAYMANOSTELLID ASTEROID REVIEW

Fig. 5. Caymanostella phorcynis sp. nov. (holotype, AMJ 18911) A = aboral view, B = oral view (R = 5.0mm).
(Paratype, NMNZ 4248), G = aboral view (arrows indicate positions of distal-lateral disc plates), D = oral
view (R = 5.0mm), E = distal-lateral disc plate (x 22), F = odontophore (x 22).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

F. W. E. ROWE 303

adambulacral plates with 1-2 furrow spines, 10-12 spines on the actinal surface;
abactinal granulation even, granules dome-shaped.

Description: The specimens range in size from R = 2.8-10.5mm,r = 2.2-8.5mm, R/r
= 1.05-1.30 (holotype: R = 5.0mm, r = 3.9, R/r = 1.28). The abactinal surface is
covered with imbricating plates almost all of which imbricate in a similar direction, with
the proximal edge of each plate overlying the distal edge of the preceding plate (Fig. 5A,
C). On the disc imbrication is less regular. However, the only plates which can be most
easily distinguished are the 5 primary interradials by virtue of their larger size than the
surrounding plates and by the location of the madreporite in interradius CD. Also
recognizable is a fan-shaped plate distal to the primary interradial plate and between it
and the proximal-most two superomarginals. A small distal-lateral disc plate is
separated either side of the overlap between the primary interradial plate and the more
distal fan-shaped plate described above (Fig. 5C, E). An additional series of 3 small
plates separates the dldp and interradial fan-shaped plate from the primary interradial
in the large specimens. The anus is surrounded by 4-5 plates.

Along the arms is a carinal row of 5-8 fan-shaped plates, either side of which is a
row of 3-6 dorsal-lateral plates (Fig. 5A, C). These latter plates alternate with the carinal
plates. Occasionally, on some arms, 2 or 3 small intercalary plates are present, which
disturbs the regularity of the dorsal arm plate arrangement. In the largest specimen
(NMNZ 4252; R = 10.5mm) 3 plates form a second dorso-lateral series, alternating
with the plates of each of the first series. The terminal plate bears a large, more or less
central perforation.

There are 8-15 superomarginal and inferomarginal plates which are elongate at
right angles to the periphery. These plates decrease in size regularly towards the arm
tips. The first superomarginal on adjacent arms is about twice as large as the next
adjacent plate. It bears, towards its centre, a gonopore. The other marginal plates are of
similar size to each other along the length of the arms. The edge of each of the
inferomarginals bears two spines.

The furrows are narrow, or at the most in smaller specimens, sub-petaloid (Fig. 5B,
D). There are 9-14 pairs of well-developed tube-feet in each ambulacrum, the number
increasing with growth. The ambulacral plates are dumb-bell-shaped.

With the exeption of the first plate, the adambulacrals are typically elongate,
obliquely perpendicular to the groove, extending to the proximal edge of the infero-
marginal plates. These plates decrease in size towards the arm tip. The adambulacral
plates bear rugose, tapering spines. The first plate bears 3-5 spines, of which the inner-
most 2, occurring slightly obliquely on the adradial edge are interpreted as furrow
spines. Up to 10-12 spines occur on each of the remaining adambulacral plates of which
2 furrow spines occur regularly on the adradial edge of the first 4 plates and irregularly
alternating with 1 furrow spine to the seventh plate, thereafter 1 furrow spine occurs on
the adradial edge of each of the adambulacral plates. The remaining spines are in two
alternating rows across each plate. The lower surface of each inferomarginal plate bears
spinelets similar in form to, but smaller than, those on the adambulacral plates (Fig. 5B,
D).

The oral plates bear 3-4 furrow spines, 5 on the largest specimen (NMNZ 4252; R
= 10.5mm) and 2-3 suboral spines.

The gonads on either side of the interradial membrane, can be seen through a more
or less triangular actinal membrane. This membrane is delimited by the oral plates, the
first 2 adambulacral plates and the innermost 2 inferomarginal plates.

The actinosome 1s large, about 4% diameter of the preserved animals.

Abactinal granulation is fine, even, with granules slightly longer than broad,
dome-shaped, slightly spaced, 9-12 per mm.

PROG. LINN. SOG. N.S.W., 111 (4), 1989

304 A CAYMANOSTELLID ASTEROID REVIEW

Both the anus and gonopores are surrounded by 7-9 granules.
Etymology: Phorcynis, daughter of the sea god Phorcus and Medusa.
Remarks: The shape and arrangement of abactinal plating easily separates C. phorcynis
from C. admiranda Belyaevy and Litvinova, which otherwise share the feature of the
gonopore piercing the proximal-most superomarginal plates.

Although they do not appear to differ significantly from Tasman Sea specimens,
the seven ‘Siboga’ specimens are identified as C. phorcynis with some hesitation due to
their very poor state of preservation. Further material in better preserved state is
required to confirm the occurrence of C. phorcynis in the Indonesian region. Otherwise
C. phorcynis is known from both sides of the Tasman Sea in depths between 736-1208m.
The ‘Siboga’ specimens were taken in deeper water at a depth of 2053m.

Belyaevostella gen. nov.

Description: A caymanostellid sea-star with body becoming invested in thick skin with
growth which includes, aborally, very thin, finely perforated, scale-like abactinal plates,
irregularly imbricating and in no apparent order; superomarginal plates squarish in
juvenile specimens, becoming elongate in adults but smaller than the elongated
inferomarginals; proximal-most and interradially adjacent superomarginals abut in
juveniles, becoming divergent in large adults (Fig. 1G); gonopores open between the
diverging, proximal-most superomarginals (Fig. 1C); aboral papulae present; actinal
membrane with perforated plate-like spicules (Fig. 7A, B); adambulacral and oral
plates as for family; distributed in East Indian region and off NE coast of Australia in
depths of 1301-1350m; habitat sunken, waterlogged wood.

Type species: Caymanostella hispida Aziz and Jangoux, 1984, by designation.

Other species included: None.

Etymology: Named for Dr. G. Belyaev who described the family. Gender feminine.
Remarks: The form and arrangement of the skeleton on the oral surface clearly shows
this genus to be a member of the family Caymanostellidae. However, the form and
arrangement of the abactinal plates, position of the gonopores, presence of spicules in
the actinal membrane and more particularly the presence of abactinal papulae, all
distinguish Belyaevostella from Caymanostella at generic level.

Belyaevostella hispida (Aziz and Jangoux)
Figs. 2C, 6A-B, 7A-B

Caymanostella hispida Aziz and Jangoux, 1984; 190, pl. 3.

Material examined: 2 specimens (holotype MNHN, ECAS 10056; paratype MNHN,
ECAS 10057), Station 220 (0°13.79"S, 188°12.73'E), 2350m, Miacassar Strait, Expe-
dition franco-indonésienne ‘Corindon.’ 1 specimen (Amsterdam Museum), Station 88
(0°34.6’N, 119°8.5’E), 1301m, Macassar Strait, ‘Siboga’ Expedition. 1 specimen
(Australian Museum J21783), Station 16 (11°41.55’S, 145°36.65"E), 2006-2053m, off
NE coast (Queensland) of Australia, collected from log, ‘Franklin’ Cruise.

Diagnosis (emended): as for genus; additionally inferomarginal plates with 3-4
elongate, acutely tipped spinelets along their margin in smaller specimens (R =
<8mm) (not observed in largest specimen, R = 13.6mm); adambulacral plates with 3-5
spines forming a simple transverse row across the plates; oral plates with 3 (not 2; Aziz
and Jangoux, 1984) furrow spines and a single suboral spine.

Remarks: The investment of the body in a thick skin, the presence of aboral papulae
and the occurrence of spicules in the actinal membrane are all new observations for this
species. The holotype (R = 8mm) and paratype (unmeasurable) are dried specimens
which are in generally poor condition. It is not surprising, therefore that the papulae

PROC. LINN. SOG. N.S.W., 111 (4), 1989

EF. W. E. ROWE

Fig. 6.

Belyaevostella hispida (Aziz and Jangoux) (AMJ 21783) A =

aboral view, B

oral view (R = 13.6mm).

PROG. LINN. SOG. N.S.W., 111 (4), 1989

306 A CAYMANOSTELLID ASTEROID REVIEW

Fig 7. Spicules from actinal membrane of Belyaevostella hispida. A = specimen R = ?<8mm (paratype,

MNHN ECAS 10057), B = specimenR = 13.6mm(AMJ21783).

were not observed by Aziz and Jangoux (1984). Also the skin investing the body is thin
and not readily apparent. However, I was able to find relatively simple spicules (Fig. 7A)
in the actinal membrane of these specimens. A third specimen, from the Macassar
Strait (Siboga Expedition) is smaller than the type specimens (R = 5.3, r = 4.5, R/r =
1.17). Although now stored in alcohol it is apparent the specimen has been dried at some
point during its storage. It therefore does not have thick investing skin cover. It does
appear to me that the investing skin develops with growth of the animal. A further
growth feature appears to be the change in shape of the interradially adjacent pair of
superomarginal plates. In the smallest (Macassar) specimen these plates are squarish
and distinctly smaller than the first pair of inferomarginal plates. However, in the type
specimens the shape of these superomarginals is already changing to becoming
elongate. In the largest specimen, which I do not hesitate to identify with the type
specimens, despite a much larger size, these superomarginals are as slender as the first
pair of inferomarginal plates, though still only about half as long. Curiously, also the two
superomarginals diverge from each other and away from the mid-interradial line.
Dissection from the ventral surface shows the gonoducts to lie in the space between the
divergent superomarginals so that the gonopores do not either notch or pierce the
adjacent marginal plates.

This distinctive caymanostellid genus interestingly shows a convergent possession
of papulae with the genera Perzbolaster Sladen, 1889, Remaster Perrier, 1894 and Anareaster
Fell and Clark, 1959 in the closely related family Korethrasteridae, but without the
development of cruciform and paxilliform abactinal plates.

PROG. LINN. SOC. N.S.W., 111 (4), 1989

F. W. E. ROWE 307
ACKNOWLEDGMENTS
I am grateful to Ms. A. M. Clark (now retired), British Museum (Natural
History), London, U.K., Dr. A. N. Baker, National Museum of New Zealand, Welling-
ton, New Zealand and Dr. M. Jangoux, Université Libre de Bruxelles, Belgium for loan
of type and other specimens and Dr. Jangoux allowed me access to some manuscript
figures he had prepared; I thank Professor D. TV. Anderson, Sydney University, N.S.W.,
Australia, for critical discussions. I thank the following staff from the Australian
Museum for their assistance: Ms P. Berents for reading and commenting on the
manuscript; Ms. E. Lynne Albertson prepared figures; Ms. Kate Lowe photographed
specimens; Mr. G. Avern assisted with S. E. M. photography; Ms. Kelly Walker typed
the manuscript.

References

Aziz, A. and JANGOUX, M., 1984. — Description de quatre novelles especes d’astérides profonds (Echinoder-
mata) de la région Indo-Malaise. Indo-Malayan Zoology 2: 187-194, pls. 1-4.

BAKER, A. N., Rowe, F. W. E. and CLARK, H. E. S., 1986. — A new class of Echinodermata trom New
Zealand. Nature Lond. 321: 862-864, figs. 1-4.

BELYAEV, G., 1974. — A new family of abyssal starfishes. Zoologicheski Zhurnal 53 (10): 1502-1508, 2 figs. (In
Russian).

BELYAEV, G. and Litvinova, N. M., 1977. — The second finding of decp-sea starfishes of the family
Caymanostcllidae. Zoologicheskit Zhurnal 56 (12): 1893-1896, 2 figs. (In Russian).

BLAKE, D. B., 1981. — A reassessment of the sea-star orders Valvatida and Spinulosida. J. Nat. Hist. 15: 375-
394, figs. 1-4.

BLAKE, D. B., 1987. — A classification and phylogeny of post-Palacozoic sea-stars (Asteroidea: Echinoder-
mata). J. Nat. Hist., 21: 481-528, figs. 1-13.

DANIELSSEN, D. C. and KOREN, J., 1884. — Yhe Norwegian North Atlantic Expedition, 1876-1878. Zoology.
Asteroidea. Christiania: 119 pp., 15 pls.

FELL, H. B. and CLark, H. E., 1959. — Anarcaster, a new genus of Asteroidea from Antarctica. Zrans. Roy.
Soc. N.Z. 87: 185-187, figs. 1-3.

HEALY, J. M., Rowr, F. W. E. and ANDERSON, D. T., 1988. — Spermatozoa and spermiogenesis in Ayloplax
(Class Concentricycloidea): a new type of spermatozoon in the Echinodermata. Zool. Scripta 17 (3):
297-310, figs. 1-63.

PERRIER, E., 1984. — Expeditions scientifique du Travailleur et du Talisman. 1 Stellerides, 431 pp., 26 pls. Paris.

RowkE, F. W. E., 1985. — On the genus Podosphaeraster A. M. Clark and Wright (Echinodermata: Asteroidca)
with description of a new species from the North Atlantic. Bull. Mus. Natn. Hist. nat. Faris 4e ser., 7,
sect. A, no. 2: 309-325, figs. 1-3, 1 pl.

RowkE, F. W. E., 1988. — Review of the extant Class Concentricycloidea and re-interpretation of the fossil
Class Cyclocystoidea. Jn BOURKE, R-(ed.). Echinoderms: Proceedings of the Sixth International Conference,
Victoria, 25-28th August, 1987. Balkema: Rotterdam: pp. 1-13, figs. 1-6.

Rowe, F. W. E., BAKER, A. N. and CLARK, H. E. S., 1988. — The morphology, development and taxonomic
status of Ayloplax Baker, Rowe and Clark (1986) (Echinodermata: Concentricycloidea), with the
description of a new species. Proc. Roy. Soc. Lond. B233: 431-459, figs. 1-72.

SLADEN, W. P., 1889. — Asteroidea. Rep. scrent. Results Voy. ‘Challenger’ (Zool.) 30: 893 pp., 117 pls.

SMITH, A. B., 1988. — ‘To group or not to group: the taxonomic position of Xploplax. In BURKE, R. (ed.)
Echinoderms: Proceedings of the Sixth International Conference, Victoria, 23-28th August, 1987. Balkema:
Rotterdam: pp. 17-23, fig. 1.

SMITH, A. B. and TRANTER, T. H., 1985. — Protremaster, anew Lower Jurassic genus of asteroid from Antarc-
tica. Geol. Mag. 122 (4): 351-359, figs. 1-6.

SPENCER, W. K. and WRIGHT, C. W.,, 1966. — Asterozoans In MOORE, R. C. (ed). Treatise on Invertebrate
Palaeontology vol. U3 (1) Echinodermata Lawrence, Kansas: University of Kansas Press, pp. U4-U107,
figs. 1-89.

PROG. LINN. SOG. N.S.W., 111 (4), 1989

re te

4 iM ney Dey

143
157

169

201

215

225

233

241

257

293

NUMBER 3

R. L. PRESSEY

Wetlands of the Lower Clarence Floodplain, northern coastal New South Wales
als de AR ES STa\¢

Wetlands of the Lower Macleay Floodplain, northern coastal New South Wales

J. W. PEMBERTON

The Ordovician — Silurian Stratigraphy of the Cudgegong — Mudgee District, New
South Wales

S. J. ROWLAND

Aspects of the History and Fishery of the Murray Cod Maccullochella peeli (Mitchell)
(Percichthyidae)

T. F. FLANNERY

Microhydromys musseri n. sp., a New Murid (Mammalia) from the Torricelli
Mountains, Papua New Guinea

NUMBER 4

A. S. STEFFE

Tidal and Diel Variations in the Abundance of Larval Fishes in Botany Bay, New
South Wales, with Emphasis on Larval Silverbiddy Gerres ovatus (Fam. Gerreidae)
and Gobies (Fam. Gobiidae)

C. M. A. KENNEDY

Redescription of Austrochthonius australis Hoff (Chthoniidae: Pseudoscorpionida)
R. A. BUCHANAN

Pied Currawongs (Strepera graculina): their Diet and Role in Weed Dispersal in
suburban Sydney, New South Wales

F. W. E. ROWE

Nine new, deep-water Species of Echinodermata from Norfolk Island and
Wanganella Bank, northeastern Tasman Sea, with a Checklist of the Echinoderm
Fauna

F. W. E. ROWE

A Review of the Family Caymanostellidae (Echinodermata: Asteroidea) with the
Description of a New Species of Caymanostella Belyaev and a New Genus

ANNEXURE to Numbers 1-4. THE LINNEAN SOCIETY OF NEW SOUTH WALES.
Record of the ANNUAL GENERAL MEETING 1987, Reports and Balance Sheets.

PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES
VOLUME 111

SOCIETY

Issued 21st December, 1989 ;
iia ee ae nS ee ea
CONTENTS:

NUMBER 1

1 M. J. LAMBERT and J. TURNER

Redistribution of Nutrients in Subtropical Rainforest Trees

is Gi. YOUNG :
New Occurrences of Culmacanthid Acantnodians (Pisces, Devonian) from
Antarctica and southeastern Australia :

25 M.R. GRAY and G. J. ANDERSON
A new Australian Species of Argyrodes Simon (Araneoidea: Theridiidae) which preys
an its:Host

31. °C. N? SMITHERS
Two new Species of Amphientomidae (Insecta: Psocoptera), the first Record of the
Family. for Australia

37. A. B. BRADKE and D. R. MURRAY
Redistribution of Amino Acids and Amides during Seedling Development in Acacia
iteaphylla F. Muell. (Fabaceae: Mimosoideae)

NUMBER 2

45 H. CAMPBELL
John Vaughan Thompson, F.L.S.

65... E.W.:GROVES and: D. T. MOORE *
A List of the Cryptogams and Gymnospermous Plant Specimens in the British
Museum (Natural History) gathered by Robert Brown in Australia 1801-5

103. J. A. ELIX and H. STREIMANN
The Lichens of Norfolk Island. 1: Introduction and the Family Parmeliaceae

123 C.M. A. KENNEDY
Conicochernes doyleae, a new Australian Species of the Chernetidae
(Pseudoscorpionida: Arachnida)

131. |. YASSINI and M. MIKULANDRA
Mckenziartia and Pectocythere (Pectocytheridae, Ostracoda, Crustacea) in Lake
Macquarie, New South Wales

Contents continued inside
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