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PROCEEDINGS
Ole Walle
LINNEAN SOCIETY
F
NEW SOUTH WALES
VOLUME 104 (Nos 457-460; for 1979-80)
Sydney The Linnean Society of New South Wales 1980
© Linnean Society of New South Wales
Contents of Proceedings Volume 104
PART 1 (No. 457) (Issued 21st July, 1980)
PEOU, S. Some Carboniferous articulate brachiopods from eastern
News Ouchi Wialesr sive rspcccnc sae cis apenas ee ee ur raw) oe aanieVn alse LARSON, H. K., and HOESE, D. F. The species of the Indo-West
Paciic genus, Galumza) (Pisces: Eleotridae) <3: 5...55.-2..45-- HARRIS, M. MCD., KING, R.J., and ELLIS, J. The eelgrass
Zostera capricornz in Illawarra Lake, New South Wales ......... HUTCHINGS, P., and RAINER, S. A key to estuarine polychaetes in
INEWESOUCIVialeS Wee seis h eo retort acts euepvias Mee os 6 oe ei ehabe BUCHANAN, R. A., and HUMPHREYS, G. S. The vegetation on
two podzols on the Hornsby Plateau, Sydney ................. BUCHANAN, R. A. The Lambert Peninsula, Ku-ring-gai Chase
National Park. Physiography and the distribution of podzols,
shrublands and swamps, with details of the swamp vegetation
ANGLES FITMENT Sy rs ctter sc citer ka Las tele aur seta )os ctio) Suisicovle Pe gessteeoeuetlau'e ceria) emistin te
PART 2 (No. 458) (Issued 21st July, 1980)
FACER, R. A., HUTTON, A. C., and Frost, D. J. Heat generation by siliceous igneous rocks of the basement and its possible influence on coal rank in the Sydney Basin, New South Wales ............ CARR, P. F., JONES, B. G., and WRIGHT, A. J. Dating of rocks from the Bungonia district, New South Wales .................... TIMMS, B. V. The benthos of the Kosciusko glacial lakes ............ McNAMARA, K. J., and PHILIP, G. M. Living Australian schizasterid SCHIMOLASMER nace Sree e tee Paces has at occ wc erctns sgh neon Mle ay as ANDERSON, D. T. Cirral activity and feeding in the lepadomorph barnacle Lepas pectinata Spengler (Cirripedia) ...............
Annexure to Parts 1 and 2. The Linnean Society of New South Wales. Record of the Annual General Meeting 1979. Reports amMGep aliameeys Mets: se reer cee ral tse ol ee otiee ove smelt @) a 6 eels oc
NUMBER* 3 (No. 459) (Issued 16th January, 1981)
SHAW, D. E., and CARTLEDGE, E. G. Sporobolomycetaceae from Indooroopilly (Australia) and from Port Moresby
(Papua New Guimea) tei ia ie er a oaiey leat nna e Wa Mani pn arebeen 161 SKILBECK, C. G. A preliminary report on the late Cainozoic geology and
fossil fauna of Bow, New South Wales ....................0- level DOMROW, R. The genus Razllzetza Trouessart in Australia (Acari:
Dermanyssid ae). jn S es oe as eer i ce licen ea ey tice ona 183
POWELL, C. MCA., FERGUSSON, C. L., and WILLIAMS, A. J. Structural relationships across the Lambian Unconformity in the Hervey Range-Parkes:area IN :SoW) ic, Lace epee Cee tee 195
WRIGHT, A. J., and FLorY, R. A. A new Early Devonian tabulate coral from the Mount Frome Limestone, near Mudgee, New South Wales 211
NUMBER 4 (No. 460) (Issued 16th January, 1981)
DOMROW, R. A new species of the ulysses group, genus Haemolaelaps Berlese(Acam: Dermanyssidae)) 2) goin a ae ee eee 22h CARR, P. F., JONES, B. G., KANSTLER, A. J., MOORE, P.S., and Cook, A. C. The geology of the Bungonia district, New South
Weales ceo Ce SORk OS ie SGN Ah Vago aeae TERR ne Ga enn Ra 229 SKINNER, S. New records of Zygnemaphyceae and Oedogoniophyceae
(Chlorophyta) from northern New South Wales .............. 245 DuGAN, K. G. Darwin and Diprotodon: The Wellington Caves
fossilsiand the law, of succession’ elias) ee ee eee 265 LEITCH, E. C. Rock units, structure and metamorphism of the Port
Macquarie Block, eastern New England Fold Belt ............. 273 LINNEAN SOCIETY OF NEW SOUTH WALES. Presidential addresses
prntedanitheiProceedmnes NOZ0719 i She eee ae eee 293 TINE 2). Ps, SU Sel ea nS TS lene nn GE ae 299
*The change from Part to Number is required by the latest guidelines governing eligibility for the Book Bounty. Issue of the record of Society business as an Annexure to Parts 1 & 2 follows the same advice. — Ed.
PROCEEDINGS of the |
LINNEAN - SOCIETY
NEW SOUTH WALES
VOLUME 104 PARTS 1&2
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OFFICERS AND COUNCIL 1979-80
President: A. RITCHIE
Vice-Presidents: LYNETTE A. MOFFAT, T. G. VALLANCE, J. T. WATER- HOUSE, B.D. WEBBY .
Honorary Treasurer: D. A. ADAMSON
Secretary: BARBARA STODDARD
Council: D. A. ADAMSON, M. ARCHER’, L. W. C. FILEWOOD?’, A. E. GREER, L. A. S. JOHNSON, HELENE A. MARTIN, P. M. MARTIN?, LYNETTE A. MOFFAT, P. MYERSCOUGH, A. RITCHIE, A. N. RODD, F. W. E. ROWE, E. J. SELBY*, C. N. SMITHERS, P. J. STANBURY, N. G. STEPHENSON’, T. G. VALLANCE, JOYCE W. VICKERY’®, J. T. WATERHOUSE, B. D. WEBBY, A.J. T. WRIGHT
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* appointed 23 May 1979 ? appointed 19 December 1979 3 appointed 21 November 1979 * resigned 23 May 1979 ° resigned 24 October 1979 ° deceased 29 May 1979
Cover motif: Fossarus sydneyenszs Hedley
Balmoral Beach, Sydney (Proc. Linn. Soc. N.S.W. 35, 1900: 89)
PROCEEDINGS of the
LINNEAN SOCIETY
NEW SOUTH WALES
VOLUME 104 PANT JI
Some Carboniferous Articulate Brachiopods from Eastern New South Wales
S. PEOU pe ee ee
PEou, S. Some Carboniferous articulate brachiopods from eastern New South Wales. Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980:1-15.
Nine fossil localities in the Carboniferous Berrico Creek Formation at Berrico- Rawdon Vale and an unnamed formation at Brownmore in eastern New South Wales, have yielded a number of new brachiopod taxa belonging to the Rhipidomella forttmuscula Zone. Among these taxa are Quadratza engeli sp. nov., Q. boonzenszs sp. nov., Productoid gen. et sp. nov., ‘Camarotoechia’ subtrigonalis sp. nov., Schizophoria subelliptica sp. nov., Podtsheremia fasciculata sp. nov., and Brachythyris cobarkensts sp. nov. They are short-ranging species and hence are suitable as zonal index fossils.
S. Peou, Department of Geology, University of Newcastle, Australia 2308; manuscript received 28 September 1978, accepted in revised form 21 February 1979.
INTRODUCTION
The Rhip:domella fortemuscula Zone is one of nine major brachiopod zones in the Carboniferous of eastern Australia (Campbell and McKellar, 1969; Campbell and Roberts, 1969; Jones et al., 1973; Roberts, 1975; Roberts et al., 1976). The fauna of this zone was first studied by Cvancara (1958) and then by Campbell and McKelvey (1972) from the Barrington District, New South Wales. McKellar (1967) and Driscoll (1960) recognized this fauna from the Yarrol Trough, Queensland.
Over the past twenty years, a large number of brachiopod genera and species have been described from the Rhzpzdomella fortimuscula Zone, many of them are short-ranging forms and clearly indicative of a late Visean age (Roberts, 1975, 1976). Recent work on rocks of this age in the Berrico Creek Formation at Berrico-Rawdon Vale (Whitford, 1971; Peou, 1977) and an unnamed formation at Brownmore (McDonald, 1972; Roberts, 1975) (Fig.1), has resulted in discrimination of several new taxa. Good stratigraphical control and short ranges suggest that these taxa may
per BERRICO-RAWDON VALE BROWNMORE BRACHIOPOD ZONES/SUBZONES CUT HILL BOORAL FORMATION FORMATION FAULKLAND FORMATION 7A 7
UNNAMED SANDSTONE BERRICO CREEK 6 FORMATION
6 CARSONVILLE FORMATION 5
WOOTTON BEDS
lL inoproductus (Balakhonia)/rawdonvalensis Marginirugus barringtonensis
UNNAMED 6 FORMATION
Rhipidomella fortimuscula
6?
pinea
FLAGSTAFF
CARBONIFEROUS
aspinosa
SANDSTONE
Fig. 1. Correlation of Carboniferous formations of the Berrico-Rawdon Vale and Brownmore districts, N.S.W. The brachiopod zones/subzones are indicated by numbers on the right hand side of columns (Modified from Roberts, 1975; Peou, 1977; and Peou and Engel, 1979).
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
2 CARBONIFEROUS ARTICULATE BRACHIOPODS
be useful for zonal identification. They include two species of Quadratza, one species each of ‘Camarotoechia’, Schizophoria, Podtsheremia and Brachythyris; one productoid described on the basis of only three specimens has been temporarily designated as a new genus and species.
The descriptions of these brachiopods are based on collections housed in the Department of Geology, University of Newcastle, New South Wales. The fossils have been obtained from nine localities in the Berrico-Rawdon Vale-Brownmore region, N.S.W. (Fig.2). Details of these localities (locality number, grid references and name of the topographic sheets) are given at the end of the paper.
SYSTEMATIC DESCRIPTION
Superfamily PRODUCTACEA Family PRODUCTELLIDAE Schuchert & LeVene 1929 Subfamily CHONOPECTINAE Muir-Wood & Cooper 1960 Genus QUADRATIA Muir-Wood & Cooper 1960 Type species: Productus hirsutiformis Walcott 1884. Remarks: The specimens described below differ from those of Quadratza from the Mississippian rocks of Oklahoma and Nevada, in having a weak ventral sulcus and dorsal fold, better developed prostrate spines on the ventral visceral disc, faintly dendritic adductor scars, broader and flat lateral ridges in the brachial valve, and a
pronounced adductor platform in the same valve. In addition, the present specimens are characterized by their bilobed cardinal process whose lobes are incised and
Gloucester,
Newcastle ryYGLOUCESTER
WARDS RIVER OUPPER CHICHESTER
SALISBURY
L5450 -OBROWNMORE
Fug. 2. Fossil locality map of the Berrico-Rawdon Vale and Brownmore districts, N.S.W.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
So JAKOG) 3
separated by a distinct median depression, instead of a knoblike process observed in Quadratia, but described by Muir-Wood and Cooper (1960, p.161) as probably incomplete. Most of these differences could be of generic significance, however it would be better to await further knowledge of the type species of Quadratza, before separate generic status is given to the present material.
Quadratza engeli sp. nov. Fig. 3, 1-7
Material: NUF 4049-4057, 4355; holotype NUF 4053, paratypes NUF 4049, 4054- 4057, all from NUL 640 (the type locality) in the Berrico Creek Formation at Rawdon Vale. Derivation of name: In honour of Associate Professor B.A. Engel, Department of Geology, University of Newcastle, N.S.W. Diagnosis: A species of Quadratza characterized by having a distinct sulcus and fold, prostrate spines on ventral visceral disc and suberect to erect spines scattered on trail, a diamond-shaped dorsal adductor platform, a conspicuous breviseptum, and a moderate cardinal process with a median depression. Description: Exterior. Shell medium-sized, transverse, widest at straight hinge; interareas bearing fine striations; ears broad and flattened. Pedicle valve with a convex visceral disc and a short trail; umbo small, overhanging hinge; valve surface bearing faint concentric growth lines and weak rugae; prostrate spines rare, suberect to erect spines not arranged in rows on trail. Brachial valve with a concave visceral disc and a steep trail; interarea narrower than that of the pedicle valve; concentric growth lines and rugae well developed. Interior. Pedicle valve having an elliptical cavity of visceral disc well demarcated from impressions of external ears; anterior adductor scars either pear-shaped or elongate, elevated, placed between shallowly depressed posterior adductor scars; a shallow furrow separating adductor scars, and bearing a median ridge; diductor scars tear- drop-shaped in outline, ridged anteriorly and smooth posteriorly, enclosing adductor scars; a large shell thickening situated posterior to muscle field; two ridges borne near elongate hinge teeth, diverging at 16° from hinge. Brachial valve with an elevated adductor platform having a median depression; adductor scars weakly dendritic, spreading from the depression to lateral slopes of platform, bounded posterolaterally by two heavy ridges supporting cardinal process and diverging at 38°-40° from hinge ; anterior adductor scars broad, embracing small triangular posterior adductor scars; breviseptum non-sulcate, arrow-headed, dividing adductor scars; cardinal process with two high lobes shallowly incised ; a weak median ridge defined in deep depression separating the two lobes, detached from breviseptum by a small alveolus; sockets deep, bounded anteriorly by short socket ridges; narrow furrows separating these ridges from lobes of cardinal process; lateral ridges broad and flat, attached to socket ridges, following hinge margin, and dying out before reaching cardinal extremities. Measurements: Length of shell: 18.5mm-25.5mm; width of shell: 36mm-44mm. Remarks: Quadratza egregia Carter (1967) from the Mississippian Chapel Limestone of central Texas, U.S.A., is similar to the present species in the presence of ventral sulcus and dorsal fold. However, Carter’s species has smaller ears, overlapping growth lamellae and small spine bases in concentric rows on the pedicle valve, small closely set flattened teeth, and a lower bifid cardinal process supported by a narrow median septum. The Mississippian species Q. hirsutiformzs (Walcott) redescribed by Muir- Wood and Cooper (1960) from Oklahoma and Nevada, differs from Q. engelz in having stronger concentric growth lines and rugae, a more transverse cavity of the ventral visceral disc, thicker but shorter ridges arising near hinge teeth, a thinner breviseptum being not arrow-headed and originating from a low smooth platform,
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
4 CARBONIFEROUS ARTICULATE BRACHIOPODS
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980.
S. PEOU 5
and weaker ridges bounding the dorsal adductor scars postero-laterally. Q. rangariensis (Campbell, 1963) from the Tournaisian rocks of the Werrie and Belvue Synclines, N.S.W., has more numerous spines on the pedicle valve, broader but shorter adductor scars and smaller diductor scars in the same valve. The brachial valve of Q. rangariensis has a non-arrow-headed breviseptum developed from a flat to rounded posterior platform, and has undivided smooth adductor scars bordered postero-laterally by two short ridges arising from the central platform.
Quadratia booniensis sp. nov. Fig.3, 8-18 Materzal: NUF 4037-4048; holotype NUF 4044, paratypes NUF 4045 and 4047, all from NUL 640 (the type locality) in the Berrico Creek Formation at Rawdon Vale.
Derivation of name: After Boonie Doon farmhouse at Rawdon Vale.
Diagnosis: A species of Quadratia having very distinct prostrate spines, suberect to erect spines in rows on ventral trail, a low dorsal platform, a thin breviseptum, and a small cardinal process with a shallow median depression.
Description: Exterior. Shell medium-sized, transverse, with a maximum width at straight hinge; interareas flat, horizontally striated; ears subtriangular, flattened. Pedicle valve with a gently convex visceral disc and a short trail; umbo small, incurved over hinge; valve smooth on trail, bearing poorly developed concentric growth lines and rugae elsewhere; 7-8 spines on one row on trail, 2-3 spines defined on hinge margin. Brachial valve with a weakly concave visceral disc and a steep trail; interarea narrower than that of the pedicle valve; valve surface bearing prominent growth lines and rugae.
Interior. Pedicle valve having an elliptical cavity of visceral disc well separated from impressions of external ears; adductor muscle field heart-shaped in outline; anterior adductor scars slightly elevated, faintly dendritic, inserted between smooth posterior adductor scars; a furrow dividing adductor scars; a weak ridge extending on floor of the furrow near posterior end of muscle field or from a shell thickening in one specimen to a short distance from anterior ends of adductor scars; diductor scars tear- drop-shaped, either smooth or faintly ridged; two distinct ridges arising in front of small and sharp hinge teeth, diverging at 14°-15° from hinge, being curved and obsolete along inner edges of ears. Brachial valve with a gently elevated adductor platform having a shallow median depression with a sharp breviseptum being obscure between adductors scars not clearly separated; a faint median ridge sitting in shallow depression between two incised lobes of cardinal process; two short but robust ridges supporting the process, diverging at 32°-34° from hinge; alveolus ill-defined; sockets deep, elongate; socket ridges developed from lobes of cardinal process by narrow furrows, lateral ridges broad and flat, running along hinge margin, fusing on ears.
Fig. 3. 1-7. Quadratia engeli sp. nov. 1. Latex cast of a pedicle valve exterior; NUF 4049, paratype, x1. 2. Latex cast of an incomplete pedicle valve exterior showing a sulcus; NUF 4051, x1.5. 3. External mold of a brachial valve; NUF 4052, x1.1. 4-5. Internal mold of two pedicle valves; NUF 4055 and 4053, both paratypes, x1.1 and x1.5 respectively. 6. Latex cast of an incomplete brachial valve interior; NUF 4047, holotype, x1.3. 7. Enlargement of NUF 4047 showing a bilobate cardinal process whose incised lobes are separated by a median depression bearing a distinct ridge, x3. All from NUL 640, Rawdon Vale.
8-18. Quadratia booniensis sp. nov. 8-9. Latex cast of two pedicle valve exteriors; NUF 4038 and 4037, both x1. 10. External mold of an incomplete brachial valve; NUF 4041, x1. 11. Latex cast of NUF 4041, x1. 12. Internal mold of a pedicle valve; NUF 4045, paratype, x1.1. 13. Latex cast of NUF 4045, x1. 14. Internal mold of a pedicle valve; NUF 4044, paratype, xl. 15. Latex cast of NUF 4044, x1. 16. Latex cast of an incomplete brachial valve interior; NUF 4057, holotype, x1.5. 17. Enlargement of NUF 4057 showing a bilobate cardinal process with incised lobes separated by a median depression, x4.2. 18. Latex cast of an incomplete brachial valve interior; NUF 4355, x1.5. All from NUL 640, Rawdon Vale.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
6 CARBONIFEROUS ARTICULATE BRACHIOPODS
Measurements: Length of shell: 18.5mm-25.5mm; width of shell: 31mm-44mm.
Remarks: Quadratza booniensts is similar to Q. engelz mainly in the size and the shape of the shell, straight hinge, striated interareas, flattened ears, tear-drop outlines of ventral diductor scars, broad lateral ridges and incised lobes of the cardinal process. However, Q. engeli differs in having distinct sulcus and fold, less developed prostrate spines on the visceral disc and scattered suberect to erect spines not arranged in rows on the trail of the pedicle valve, elongate ventral adductor scars, a better developed dorsal adductor platform, a thicker and arrow-headed breviseptum, a stronger cardinal process supported on two heavier ridges, and a deeper median depression separating the two lobes of the process. Q. hzrsutzformzs (Walcott) redescribed by Muir-Wood and Cooper (1960) possesses a more pronounced concentric ornament, a more transverse cavity of ventral visceral disc, deeply impressed ventral adductor scars, a better developed alveolus, a shorter breviseptum springing from a smooth platform, and weaker ridges supporting the cardinal process. Q. egregza Carter (1967) is characterized by having distinct sulcus and fold, developed overlapping growth lamellae, several small spine bases on each side of ventral umbo close to posterior margin, closely set flattened teeth, and a narrow median septum supporting the cardinal process. The only other comparable species, Q. rangariensts (Campbell, 1963), has strongly developed spines on the pedicle valve, suboval to subtriangular ventral adductor scars being slightly embraced by the small diductor scars, a broad posterior dorsal platform from which arises a more distinct breviseptum, and weaker and shorter ridges bounding dorsal adductor scars.
Family DICTYOCLOSTIDAE Stehli 1954 Subfamily DICTYOCLOSTINAE Stehli 1954 PRODUCTOID gen. et sp. nov. Fig. 4, 1-6 Materzal: NUF 4021-4023, all from NUL 545 in the unnamed formation at Brownmore.
Description: Exterior. Shell large, concavo-convex; hinge line short; interareas concave; ornament of costae forming a reticulation with weak concentric ridges on umbonal region, broad concentric lamellae (on brachial valve only) , growth lines and spines. Pedicle valve with a small umbo, a shallow sulcus, a broad visceral disc and a long trail; costae subrounded to flat, branching, curved posteriorly on steep lateral slopes, and wrapping high knoblike structures on small ears; spines suberect to erect, big and long. Brachial valve with a short umbo, a low fold, and small ears bearing deep depressions for the reception of knoblike structures on ventral ears; costae
Fig. 4. 1-6. Productoid gen. et sp. nov. 1. Latex cast of a pedicle valve exterior; note the distinct knoblike structure on one ear; NUF 4021, figured specimen, x0.8. 2. Latex cast of an incomplete pedicle valve exterior; NUF 4023, x1.6. 3. Latex cast of a cardinal area and umbonal regions of pedicle valve and brachial valve exteriors; NUF 4023, x2. 4-5. External mold of a brachial valve showing the broad visceral disc (4) and trail (5) ; NUF 4023, x0.9 and x0.8 respectively. 6. External mold of a brachial valve; note the distinct reticulate and lamellose ornament; NUF 4022, figured specimen, x0.9. All from NUL 545, Brownmore.
7-17. ‘Camarotoechia’ subtrigonalis sp.nov. 7. Latex cast of a pedicle valve exterior; NUF 4059, x3. 8.Latex cast of a brachial valve exterior; NUF 4058, x2.6. 9-10. Internal mold of two pedicle valves; NUF 4062 and 4061, both paratypes, x2.8 and x3.5 respectively. 11. Latex cast of NUF 4061, x4. 12. Internal mold of a brachial valve; NUF 4066, holotype, x4.5. 13. Latex cast of NUF 4066, x3.5. 14-15. Internal mold of a brachial vaive and latex cast of same; NUF 4065, paratype, x4.5 and x4.9 respectively. 16-17. Internal mold of two brachial valves; NUF 4067, paratype and NUF 4068, x4.2 and x5 respectively. All from NUL 723,
except NUF 4059 and 4062 from NUL 882, Rawdon Vale.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
S. PEOU
(1979) 1980
Proc. Linn. Soc. N.S.W., 104 (1)
8 CARBONIFEROUS ARTICULATE BRACHIOPODS
rounded, increasing by intercalation; big spines rare, faint spines on long trail. Internal structures of both valves unknown.
Measurements: Length of shell: 43mm-51mm; width of shell: 52mm-56mm. Remarks: Only three specimens are at present available for the description. The brachial valve exhibits three different ornamental zones: (a) a reticulate zone occupying the umbonal region and being produced by the intersection between concentric ridges and radial costae; (b) a lamellose zone situated between the other two, covering over 80% of valve surface, and bearing concentric lamellae with growth lines interrupting the radial costae, and rare spines; and (c) a narrow spinose zone on trail, bearing fine spines.
Determination of the generic affinities of the species described above is difficult, due to the lack of specimens detailing the internal structures. This form is referred to the subfamily Dictyoclostinae only on the basis of its reticulate umbonal region and costate trail. A precise assignment must await detailed study of further material. The external ornamentation of the brachial valve, especially the broadly developed concentric lamellae traversing the radial costae, and the knoblike structures on the ears of the pedicle valve are the most distinctive features which could separate the present genus from all other productid genera.
Superfamily RHYNCHONELLACEA Family CAMAROTOECHIIDAE Schuchert & LeVene 1929 Subfamily CAMAROTOECHIINAE Schuchert & LeVene 1929 Genus CAMAROTOECHIA Hall & Clarke 1893 Type species: Atrypa congregata Conrad 1841.
Remarks: Numerous workers have recorded species of Camarotoechia from the Devonian and Carboniferous of Australia. The genus has yet to be unequivocally recorded from the southern hemisphere, or for that matter from Carboniferous rocks, so the following species, a more strongly-ribbed form than the type species C. congregata (Conrad, 1841), is referred to the genus as a procedural gambit until such time as the systematics of Australian Devonian and Carboniferous rhynchonellaceans are better known.
‘Camarotoechia’ subtrigonalis sp. nov. Fig. 4, 7-17 Material: NUF 4058-4070; holotype NUF 4066, paratypes NUF 4061, 4062, 4065 and 4067, from NUL 723 (the type locality) and 882 in the Berrico Creek Formation at Rawdon Vale. Derivation of name: subtrigonalis refers to the not completely trigonal shell outline.
Diagnosis: A species of ‘Camarotoechia’ with a small and plicate shell being subtrigonal in outline, a feebly uniplicate anterior commissure, a V-shaped dorsal septalium supported by a short median septum, and an unsplit to split hinge plate bearing two short crura.
Description: Exterior. Shell unequally biconvex, generally wider than long, ornamented with coarse rounded costae. Pedicle valve moderately convex; umbo slightly incurved; sulcus distinct, shallow, commencing in front of umbo, having 3 to 5 plicae; lateral slopes not steep, 4-6 plicae on each slope; concentric growth lines poorly developed. Brachial valve more convex than pedicle valve; umbo strongly incurved ; fold low, having 4 plicae; lateral slopes steep, each slope bearing 5-6 plicae.
Interior. Pedicle valve with ill-defined adductor scars, smooth elongate diductor scars
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
S. PEOU 9
tapering posteriorly and sitting between two short and sharp dental lamellae; diverging angle of the lamellae 28°-34°; teeth strong, supported on dental lamellae. In brachial valve, adductor scars smooth, separated by a sharp median septum extending about one-half total length of valve; septalium shallow; sockets deep and elongate, bounded posteriorly by outer edges of hinge plate.
Measurements: Length of shell: 6mm-12.5mm; width of shell: 7mm-15mm.
Remarks: ‘Camarotoechia’ subtrigonalis is similar to C. sp. Campbell (1957) and C. sp. A. Roberts (1963) respectively from Babbinboon and Lewinsbrook, N.S.W., particularly in the shape and size of the shell, and the number of plicae in the ventral sulcus. C. sp. differs from the described species in the possession of longer dental lamellae and developed rays on crura. C. sp. A. has a more transverse shell, fewer plicae on dorsal lateral slopes, a shorter median septum and narrower sockets in the brachial valve. C. sp. B_ Roberts (1965) from Trevallyn, N.S.W.., is characterized by a larger, subequally biconvex shell ornamented with angular plicae on the lateral slopes, a globular to rounded brachial valve, and a shorter dorsal median septum. In addition, C. amnica and C. septima Veevers (1959) respectively from the Carnarvon and Bonaparte Gulf Basins, Western Australia, have a pentagonal shell, a deeper ventral sulcus with fewer plicae, and a weaker dorsal median septum.
Superfamily ENTELETACEA Family ENTELETIDAE Waagen 1884 Subfamily SCHIZOPHORIINAE Schuchert & LeVene 1929 Genus SCHIZOPHORIA King 1850 Type species: Conchylzolithus (Anomites) resupinatus Martin 1809.
Schizophoria subelliptica sp. nov. Fig.5, 1-7
Material: NUF 3825-3847; holotype NUF 3839, paratypes NUF 3829-3831, 3840- 3842, all from NUL 514 (the type locality) in the Berrico Creek Formation at Rawdon Vale.
Derivation of name: subelliptica refers to the not completely elliptical shell outline.
Diagnosis: A species of Schizophorza with a subelliptical shell outline, ventral adductor scars on a high elevation and deeply depressed diductor scars, strong pallial markings, and two subparallel main pallial trunks arising at anterior ends of ventral adductor scars.
Description: Exterior. Shell transverse, widest at about midlength; cardinal extremities well rounded; hinge about two-thirds to four-fifths maximum width of shell; capillae rounded, increasing by both intercalation and bifurcation, numbering 40 per 10mm on anterior median portion of shell; growth lines developed, spine bases not observed. Pedicle valve convex on umbonal region, concave anteriorly; beak small and short; lateral slopes steep; sulcus shallow, not reaching umbo; interarea broad; delthyrium open, triangular, as wide as high. Brachial valve strongly convex; lateral slopes very steep; fold indistinct; cardinal extremities gently concave; interarea narrow, bearing faint horizontal striations; notothyrium slightly wider than high.
Interior. Pedicle valve with robust dental plates bordering muscle field laterally and diverging at 58°-64°; teeth strong; diductor scars tapering posteriorly, either smooth or weakly striated, adductor scars smooth, narrowly elongate, sitting on flanks of a heavy longitudinal elevation; a shallow furrow defined on the elevation in some specimens, dividing adductor scars; vascula genitalia on anterior and lateral sides of
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10 CARBONIFEROUS ARTICULATE BRACHIOPODS
muscle field. In brachial valve, brachiophores projecting from either side of notothyrial cavity, diverging at 62°-76°; sockets elongate, with a subrounded floor bearing transverse grooves which extend to distinct fulcral plates; cardinal process consisting of a high median lobe and two lateral lobes separated by deep furrows; lamellose myophores developed in the process; anterior adductor scars shallowly depressed, separated from small posterior adductor scars by two low broad ridges; adductor scars finely striated; a large swelling concave posteriorly, located between adductor scars, bearing a rounded median ridge; postero-lateral sides of muscle field pustulose; two main parallel trunks branching anteriorly, arising either from anterior ends of posterior adductor scars or from the broad ridges separating the anterior and posterior adductor scars; another two subparallel trunks borne at anterior ends of anterior adductor scars; secondary pallial markings prominent in both valves.
Measurements: Length of shell: 19.5mm-27.5mm; width of shell: 26.5mm-33mm.
Remarks: Schizophoria subelliptica resembles S. verulamenszs Cvancara (1958) from Barrington, N.S.W., in the shape of the shell, and subparallel pallial trunks defined in the pedicle valve. It is distinguished from Cvancara’s species by its more numerous capillae, smaller diverging angles of dental plates and brachiophores, narrower ventral adductor scars, better developed secondary pallial markings, sockets and fulcral plates bearing transverse grooves, and trilobate cardinal process containing lamellose myophores. S. sp. cf. S. resupinata (Martin) described by Roberts (1971) from Bonaparte Gulf Basin, northwestern Australia, differs from the present species in having a shorter hinge, a broader ventral muscle field, a larger dorsal median swelling, a larger diverging angle of the brachiophores, and a single to multi-lobed cardinal process. S. resupznata (Martin) described by Sarycheva et al. (1963) and S. altaica Besnossova (1968) respectively from Kuznetsk Basin and eastern Kazakhstan, U.S.S.R., have a longer hinge and a less convex brachial valve. In addition, S. resupinata has fewer radial ribs, a weak dorsal median furrow, broader ventral muscle scars and stronger dorsal brachiophores; S. altazca is larger and has a rounded shell, an almost flat pedicle valve and no developed ventral sulcus and dorsal fold. Two species, S. antzqua Solle and S. striatula (Schlotheim), described by Pocock (1966) from the Devonian rocks of Germany, are characterized by having an elliptical to quadrate shell with a better developed radial ornament, an angular dorsal median septum, and fewer main pallial trunks in the brachial valve. Also, S. antzqua is smaller and possesses broader ventral muscle scars and oval dental sockets; S. strzatula exhibits stronger ventral sulcus and dorsal fold, and better developed myophores.
Fig. 5. 1-7. Schizophoria subelliptica sp.nov. 1. Latex cast of a pedicle exterior; NUF 3825, x1.6. 2. Latex cast of a brachial valve exterior; NUF 3827, x2.2. 3. Internal mold of a pedicle valve; NUF 3832, x1.2. 4. Latex cast of NUF 3832, x1.3. 5. Internal mold of a pedicle valve; NUF 3831, paratype, x1. 6. Internal mold of a brachial valve; NUF 3839, holotype, x1.1. 7. Latex cast of NUF 3839, x1.2. All from NUL 514, Rawdon Vale.
8-14. Podtsheremia fasciculata sp. nov. 8. Latex cast of a pedicle valve exterior; NUF 4152, paratype, x2. 9. Latex cast of a brachial valve exterior with an apical portion of pedicle valve exterior; NUF 4151, paratype, x1.7. 10. Latex cast of a brachial valve exterior; NUF 4155, x2. 11. Internal mold of a pedicle valve; NUF 4157, x1.7. 12. Latex cast of NUF 4157, x1.9. 13. Internal mold of a brachial valve and a posterior portion of pedicle valve; NUF 4341, holotype, x1.5. 14. Latex cast of NUF 4341, x1.6. All from NUL 628, Rawdon Vale, except NUF 4155 from NUL 861, Berrico.
15-21. Brachythyres cobarkensis sp. nov. 15. Latex cast of a pedicle valve exterior; NUF 4225, x2.1. 16. Latex cast of a brachial valve exterior; NUF 4226, x1.7. 17-19. Internal mold of three pedicle valves; NUF 4231, paratype, x1; NUF 4234, holotype, x1; and NUF 4233A, x1.1. 20. Internal mold of a brachial valve ; NUF 4243, paratype, x1.1. 21. Internal mold of a brachial valve with a posterior portion of pedicle valve ; NUF 4232B, paratype, x1. All from NUL 517, except NUF 4226, 4243 and 4232 B from NUL 515, Rawdon Vale.
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
11
S. PEOU
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
12 CARBONIFEROUS ARTICULATE BRACHIOPODS
Superfamily SPIRIFERACEA Family SPIRIFERIDAE King 1846 Genus PODTSHEREMIA Kalashnikov 1966 Type species: Podtsheremza prima Kalashnikov 1966 Podtsheremza fasciculata sp. nov. Fig.5, 8-14 Material: NUF 4341, 4151-4152, 4155, 4157; holotype NUF 4341, paratypes NUF 4151, 4152, from NUL 628 (the type locality) in the Berrico Creek Formation at Rawdon Vale, and NUL 861 in the same formation at Berrico. Derivation of name: fasciculata refers to the fasciculate costae on shell.
Diagnosis: A species of Podtsheremza characterized by its biconvex shell with rounded costae frequently branching and forming fascicles on lateral slopes, ventral sinus with a simple median costae and branching costae, dorsal fold bearing a median furrow, and rows of denticles along hinge.
Description: Exterior. Shell triangular in outline, transverse, mucronate, widest at hinge; mucros flat to slightly concave, well developed in small specimens; radial costae, fine lirae and concentric growth lines forming shell ornament; 20 to 23 fasciculate costae on each steep lateral slope, 6 to 7 simple costae on postero-lateral extremities, fascicles of 3 to 4 ribs on either side of ventral sinus and dorsal fold. Pedicle valve with a moderate and incurved umbo, a flat to weakly concave apsacline interarea ornamented with faint horizontal and vertical striations: delthyrium open, with an angle of 68°; sinus shallow, reaching umbo;,sinal angle about 19°. Brachial valve with a short umbo overhanging a narrow interarea; fold costate, conspicuous.
Intertor. Pedicle valve with high and sharp dental lamellae, supporting elongate teeth; denticles arranged in rows on either side of base of a shallow delthyrial cavity, oriented perpendicular to hinge; adminicula short and sharp, diverging at 10°-12°; adductor scars narrowly elongate, finely striated, and divided by a weak myophragm; diductor scars also elongate, smooth but faintly striated posteriorly; two ridges ill- defined between the two muscle scars; vascula genitalia observed on lateral sides of muscle field. Brachial valve with shallow sockets, distinct inner socket ridges diverging at 82° and terminating short crura; cardinal process supported on these ridges, containing up to 18 tiny vertical plates; adductor scars of two pairs: lateral pairs subtriangular; median pairs quadrangular, well impressed, divided by a fine median ridge; the two pairs being smooth, separated by two ridges corresponding to furrows which border external fold.
Measurements: Length of shell: 12mm-17mm; width of shell: 17mm-27mm.
Remarks: When compared with the type species, Podtsheremia prima Kalashnikov (1966) from northern Urals, U.S.S.R., P. fasciculata has a more transverse shell with a hinge reaching its maximum width and mucronate cardinal extremities. P.? humilicostata and P.? thomast Roberts (1971) from the Bonaparte Gulf Basin, northwestern Australia, are distinguished from the present species by their fewer costae on lateral slopes and larger angle of adminicula. In addition, P.? humilzcostata has a smaller delthyrial angle, but a larger sinal angle; P. ? thomasz has a larger angle of inner socket ridges and more numerous vertical plates in the cardinal process.
Family BRACHYTHYRIDIDAE Fredericks 1919 (1924) Genus BRACHYTHYRIS McCoy 1844 Type species: Spirifera ovalis Phillips 1836.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
S. PEOU 13
Brachythyris cobarkensis sp. nov. Fig.5, 15-21
Material: NUF 4225-4243; holotype NUF 4234, paratypes NUF 4231, 4232 and 4243, from NUL 517 (the type locality) and 515 in the Berrico Creek Formation at Rawdon Vale.
Derivation of name; After Cobark River in Rawdon Vale.
Diagnosis: A species of Brachythyris with a parasulcate anterior commissure, broad ventral muscle field from which radiate strongly developed pallial markings, robust dental ridges supporting heavy teeth, large socket plates attached to subtriangular cardinal process.
Description: Exterior. Shell unequally biconvex, with 10-12 broad and flat plicae on each lateral slope and developed concentric growth lines; plicae bifurcating, simple on cardinal extremities. Pedicle valve strongly convex on umbonal region; umbo erect, small, sharply pointed; cardinal areas broad and concave; delthyrium open, triangular, wider than high; delthyrial angle 63°; sinus shallow, reaching umbo, costae in sinus not observed, sinal angle about 9°. Brachial valve with a small and short umbo, a distinct fold bearing a weak median furrow.
Interior. Pedicle valve thickened on umbonal region; muscle scars well impressed, variable in outline, occupying about one-third valve surface; adductor scars narrowly elongate, ridged, divided posteriorly by a distinct myophragm; diductor scars broad, fusiform, having a deep posterior median furrow. In brachial valve, socket plates enclosing elongate sockets, adductor scars finely striated and divided by a weak median ridge; cardinal process having up to 32 branching, bifurcating and simple thin vertical plates; pallial markings weakly developed.
Measurements: Length of shell: 24mm-29mm; width of shell: 23mm-28mm.
Remarks: Brachythyris solida Campbell (1963) from the Belvue Syncline, N.S.W.., is similar to B. cobarkensis in the width/length ratio of shell, and the strong convexity of pedicle valve towards the umbo. However, it differs in having more numerous but narrower costae on the lateral slopes, and a subtriangular cardinal process containing fewer vertical plates. B. pseudovalis Campbell (1957) and B. elliptica Roberts (1963) respectively from Babbinboon and Lewinsbrook, N.S.W., are larger and have better developed costae on the lateral slopes, weaker sinus and fold, a larger sinal angle, and more numerous vertical plates in the cardinal process. B. planulata Roberts (1971) from the Bonaparte Gulf Basin, northwestern Australia, is characterized by its more transverse shell, uniplicate commissure, smaller delthyrial angle, larger sinal angle, and fewer vertical plates in the cardinal process. The species described by Carter (1967) as B. chouteauensts (Weller) and B. gzrty: (Branson) from the Mississippian Chapel Limestone of central Texas, exhibit better developed costae being usually simple, and lack a ventral myophragm. In addition, B. chouteauensis has a uniplicate anterior commissure, costate ventral sulcus and dorsal fold, smaller teeth and no developed dental ridges; B. gzrtyz has a subequally biconvex shell, narrower dental ridges and no developed dorsal median ridge. B. peculzar’s (Shumard) described by Weller (1914) from the Mississippian Chouteau Limestone at Mississippi Valley Basin, is smaller and possesses narrower cardinal areas, a rounded ridge dividing the lateral slopes into two regions, and fewer plicae being only simple and rounded.
LOCALITIES OF FIGURED SPECIMENS Locality number Grid references
NUL 514 738 453 Cobark 1:31680 Sheet
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
14 CARBONIFEROUS ARTICULATE BRACHIOPODS
NUL 515 740 455 Cobark 1:31680 Sheet NUL 517 736 490 Cobark 1: 31680 Sheet NUL 545 625 077 Dungog 1: 63360 Sheet NUL 628 732 499 Cobark 1:31680 Sheet NUL 640 708 462 Cobark 1: 31680 Sheet NUL 723 737 553 Cobark 1:31680 Sheet NUL 861 851 303 Gloucester 1 :63360 Sheet NUL 882 707 463 Cobark 1:31680 Sheet
NU = University of Newcastle
ACKNOWLEDGEMENTS
I am indebted to Professor B. Nashar for her continuing support, and to Associate Professor B. A. Engel for his helpful advice and critical reading of the manuscript. Facilities provided by the Department of Geology of the University of Newcastle, N.S.W., are also acknowledged.
References
BrsnossovA, G. A., 1968 — Schizophoriidae. In Sarycheva, T. G., (ed.), Brakhiopody verkhnego Paleozoya vostochnogo Kazakhstana [Brachiopods from the Upper Palaeozoic of eastern Kazakhstan]. Trudy paleont. Inst. 121:53-54 (Russian).
CAMPBELL, K. S. W., 1957. — A lower Carboniferous brachiopod-coral fauna from New South Wales. J. Paleont. 36 (1) :34-98.
———, and ENGEL, B. A., 1963. — The fauna of the Tournaisian Tulcumba Sandstone and its members in the Werrie and Belvue Synclines, N.S.W.J. geol. Soc. Aust: 10(1) :55-122.
——, and McKE iar, R. G., 1969. — Eastern Australian Carboniferous invertebrates: sequence and affinities. Pp.77-119 zn Campbell, K. S. W., (ed.), Stratigraphy and Palaeontology. Canberra: Aust. Nat. Univ. Press.
———, and McKeELvey, B. C., 1972. — The geology of the Barrington district, N.S.W. Pacific Geol. 5: 7- 43,
———, and RopsertTs, J., 1969. — Faunal sequence and overseas correlation (Carboniferous). In Packham, G.H., (ed.), The geology of New South Wales. J. geol. Soc. Aust. 16 (1) :261-264.
CARTER, J. L., 1967. — Mississippian brachiopods from the Chapel Limestone of Central Texas. Bull. Amer. Paleont. 23 (238) : 253-449.
ConrabD, T. A., 1841. — On the paleontology of the State of New York. N.Y. State Geol. Surv. 5th Ann. Rept: 25-27.
Cvancara, A. M., 1958. — Invertebrate fossils from the lower Carboniferous of New South Wales. /. Paleont. 32(5) : 846-888. ;
DRIscott, E. G., 1960. — Geology of the Mundubbera district. Pap. Dep. Geol. Univ. Qd 5(5) : 27p.
Jones, P. J., et al., 1973. — Correlation chart for the Carboniferous System of Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 156A: v+ 40p.
KALASHNIKOV, N. V., 1966. — Brakhiopody Nizhnego Karbona Verkhnei Pechory na Severnom Urale — Stratigrafia i Paleontologii severo-vostoka evropeiskoi chasti SSSR [Lower Carboniferous brachiopods of the upper Pechora in the northern Urals. Jn Stratigraphy and paleontology of the northeast European regions of the U.S.S.R.]. Akad. Nauk. SSSR, Komi Filial, Inst. Geol. 28-61 (Russian) .
McDona_p, L. K., 1972. — The geology of the Brownmore district, New South Wales. Kensington: University of New South Wales, B.Sc. thesis, unpubl. ,
McKELLAR, R. G., 1967. — The geology of the Cannindah Creek area, Monto district, Queensland. Publs. geol. Surv. Qd 331: 38p.
Muir-Woop, H., and Cooper, G. A., 1960. — Morphology, classification and life habits of the Productoidea (Brachiopoda). Mem. geol. Soc. Amer. 81: 447p.
Peou, S., 1977. — Stratigraphy, palaeoecology and taxonomy of the Rhzpzdomella fortemuscula and Balakhonia rawdonvalensis faunas in the region north of Newcastle, New South Wales. Newcastle: University of Newcastle, Ph.D. thesis, unpubl.
——, and EncEL, B. A., 1979. — A Carboniferous fauna from Rawdon Vale, New South Wales. Alcheringa 3 (2) : 141-157.
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Sp. OL) 15
Pocock, Y. P., 1966. — Devonian schizophoriid brachiopods from western Europe. Palaeont. 9 (3) :381- 412.
RoseErTs, J., 1963. — A lower Carboniferous fauna from Lewinsbrook, New South Wales. J. Proc. R. Soc. N.S.W. 97: 1-31.
——, 1965. — A lower Carboniferous fauna from Trevallyn, New South Wales. Palaeont. 8(1) : 54-81.
——, 1971. — Devonian and Carboniferous brachiopods from the Bonaparte Gulf Basin, northwestern Australia. Bull. Bur. Mener. Resour. Geol. Geophys. Aust. 122: ix + 319p.
——, 1975. — Early Carboniferous brachiopod zones of eastern Australia. J. geol. Soc. Aust. 22(1) : 1-31.
——, 1976. — Carboniferous chonetacean and productacean brachiopods from eastern Australia.
Palaeont. 19(1) : 17-77. , et al., 1976. — Late Carboniferous marine invertebrate zones of eastern Australia. Alcheringa 1 (2) : 197-225.
SARYCHEVA, T. G., et al., 1963. — Brakhiopody i paleogeografia Karbona Kuznetskoi kotloviny [ Carboniferous brachiopods and palaeogeography of the Kuznetsk Basin]. Trudy paleont. Inst. 95: 3-406 (Russian) .
VEEVERS, J., 1959. — Devonian and Carboniferous brachiopods from northwestern Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust. 45: 220p.
WELLER, S., 1914. — The Mississippian Brachiopoda of the Mississippi Valley Basin. Monogr. geol. Surv. Ill. 1: 1-508.
WHITFORD, D. J., 1971. — The geology of the Stratford-Berrico district, New South Wales. Newcastle: University of Newcastle, B.Sc. thesis, unpubl.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
The Species of the Indo-West Pacific Genus Calumza (Pisces: Eleotridae)
H. K. LARSON and D. F. HOESE
(Communicated by J. R. PAXTON)
Larson, H. K., & Hoess, D. F. The species of the Indo-west Pacific genus Calumia (Pisces: Eleotridae) . Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980:17-22.
A brief diagnosis of the marine fish genus Calumza is given. Eleotrzs godeffroy: Ginther is placed in Calumza as a senior synonym of the type species C. bzocellata Smith. Calumia profunda is described as a new species from the Solomon Islands and the New Hebrides. The species of Calumza are among the smallest-sized eleotrids known, and Calumia is one of the few eleotrid genera found on coral reefs.
It is suggested that the genus Allomicrodesmus be transferred to the family Eleotridae.
H. K. Larson and D. F. Hoese, Department of Ichthyology, The Australian Museum, P.O. Box A285, Sydney South, Australia 2000; manuscript received 15 June 1978, accepted in revised form 21 February 1979.
INTRODUCTION
The family Eleotridae contains about forty genera of which only four, Allomicrodesmus, Xenisthmus, Calumia and an undescribed genus related to Xenisthmus, are confined to coral reefs of the Indo-west Pacific. The remaining genera are inhabitants of fresh or brackish water. Allomzcrodesmus has been included in the family Microdesmidae (Schultz 1966), but it is apparent that the genus belongs elsewhere (Dawson, pers. comm.). Re-examination of the type and an additional specimen of Allomizcrodesmus dorotheae from the Great Barrier Reef shows that the species has six branchiostegal rays characteristic of eleotrids; provisionally therefore we place Allomicrodesmus in the family Eleotridae. The undescribed genus and Allomicrodesmus are both monotypic and Xenzsthmus contains about 10 species. Previously only a single species of Calumza was recognized. Examination of recently- collected material has indicated that a second undescribed species of Calumza exists. While examining types of gobioid fishes, it was discovered that C. bzocellata is a senior synonym of Eleotris godeffroyi Gunther (1877). Inaccuracies in the original description have prevented adequate identification of the species.
All of the marine eleotrid species are relatively small sized, ranging from 15 to 35 mm SL as adults. Few specimens have been collected and little is known of their distribution. For example, the species described here is known from only three specimens.
While most marine eleotrids are highly specialized forms, Calumza maintains a typical eleotrid physiognomy, with a broad scaled interorbital and a short robust body. The other genera are elongate with a protrusible lower jaw, presumably for burrowing in sand.
METHODS
Counts and measurements follow those given by Hubbs and Lagler (1958), except as listed below. The lateral scale count is the number of scale rows from the upper pectoral base to the end of the hypural plate. The transverse scale count (TRB) is taken from the anal base upward and backward to the base of the second dorsal fin. Colour notes of C. godeffroy: are based on freshly-collected specimens from the Great
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
18 SPECIES OF CALUMIA
Barrier Reef. Specimens studied are deposited in the following institutions: Australian Museum, Sydney, AMS; Bernice P. Bishop Museum, Honolulu, BPBM; British Museum (Natural History), London, BMNH; and J.L.B. Smith Institute of Ichthyology, Rhodes University, Grahamstown, RUSI. The osteology was studied from a single cleared and stained specimen of C. godeffroyy.
SYSTEMATIC DESCRIPTION Calumia Smith 1958
Calumza Smith 1958: 148 (type species: C. bzocellata Smith 1958, by original designation) .
Calumia is readily distinguished from other eleotrids by the following combination of characters. Head depressed. Top of head, cheeks and opercles scaled ; an enlarged scale between eyes. Anterior nostril elongate, positioned on snout just behind upper lip. Gill opening broad, extending forward to below preoperculum or eye. Pectoral rays unbranched. No lateral line head pores. Sensory papillae sparse, in characteristic rows (Fig. 3). Branchiostegal rays 6. First dorsal rays VI. Second dorsal rays I, 6-8. Anal rays I, 6-7. Segmented caudal rays typically 15. Lateral scale rows 21- 25. Adults small sized, reaching 15 to 20 mm SL. Vertebrae 10 + 15. Caudal skeleton with two epurals. First dorsal pterygiophore inserted after third neural spine; second and third between fourth and fifth neural spines; fourth and fifth between fifth and sixth neural spines; sixth between sixth and seventh neural spine; pterygiophore from spine in second dorsal fin between seventh and eighth neural spine, without any interneural gap. Dorsal postcleithrum present.
In genera] appearance Calumza is most similar to Ophzocara. Calumiza differs from that genus in lacking head pores, the arrangement of sensory papillae, the simple pectoral rays and the small size of adults.
KEY TO SPECIES 1. Mouth short, reaching to below anterior margin of eye. Gill opening extends forward to below posterior preopercular margin. Caudal fin with two enlarged black spots, one at upper caudal base and one on lower caudal base. Tongue tip pointed or slightly rounded. Pectoral rays 16-17. Gill rakers on outer face of lower part of first arch 6; short and stubby. Pelvic rays branched. Third and fourth dorsal spines longer than other spines ............... C. godeffroy: (Gunther) 2. Mouth longer, reaching to below posterior half of pupil. Gill opening broader, extending forward to below middle or posterior end of pupil. Caudal fin without enlarged black spots: Tongue tip bilobed. Pectoral rays 14-15. Elongate and pointed gill rakers on outer face of first arch 13-14. Pelvic rays unbranched. Second dorsallspime/lomgese, ais cia ences Mattes ec aia eee C. profunda n.sp.
Calumia godeffroy: (Gunther) Fig. 1 Eleotris godeffroy: Gunther 1877: 188, p. 122, fig. B, (Raiatea, Tahiti) . Calumia biocellata Smith 1958: 148, p. II, K and fig. 8 (Zanzibar).
Counts of the holotype of Eleotris godeffroy: are indicated with an asterisk and counts of the holotype of C. bzocellata are indicated with a plus. First dorsal rays VI (in 6)* +. Second dorsal rays I, 6(3)* +; I, 7(3). Anal rays I, 6 (3) +; I, 7 (3)*. Pectoral rays 16 (2); 17 (4)*+. Lateral scale count 21 (1); 22 (4)*+; 23 (1). Transverse backward count 7 (3); 8 (2) +. Segmented caudal rays 15 (6) * +. Lower gill rakers on outer face of first arch 6 (4) +. Predorsal scales 7 (4) *; 8 (1); 9 (1) +.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
H.K. LARSON AND D. F. HOESE 19
Fig. 1. Calumia godeffroy:; holotype of C. bzocellatus (after Smith, 1958).
Smith (1958) has described the colouration of this species, under the name Calumza brocellata, in detail. Across the nape is a transverse dark band which begins above the end of the opercle and may be intensified as a distinct black blotch on either side of the nape. The body has five greyish black transverse bands, and a trace of a sixth incomplete band at the base of the caudal rays. At the base of each pelvic fin is a small black spot. The vertical fins are black. The two caudal spots are darkest at the caudal base and tend to fade posteriorly into the dusky anterior half of the caudal. In fresh material from the Great Barrier Reef the posterior third of the second dorsal and anal fins and the posterior half of the caudal fin are yellowish orange.
This species is distinctive in the features given in the key and has been adequately described by Smith (1958). Although the original description of Eleotris godeffroyz is brief, the species is well figured. Unfortunately, Gunther (1877) did not mention or figure the two black caudal spots, but only indicated that the vertical fins are black. Examination of the holotype shows the spots and the five transverse body bands.
Calumia godeffroy? is a small-sized species, reaching a maximum size of 20 mm SL. The species is known from several localities in the western Indian Ocean (Smith, 1958), Christmas Island just south of Java (Allen, 1979) the Great Barrier Reef, Australia and Tahiti, and is undoubtedly widespread in the Indo-west Pacific. Smith (1958) reported collecting the species at low tide in muddy or weedy areas. On the outer islands of the Great Barrier Reef, the species was collected among coral and rubble at depths of 7 to 30 m.
' Material Examined: BMNH 1877.4.26.8, a 20.5 mm female, holotype of Eleotrzs godeffroyz, Tahiti. RUSI 217, 1 (29), holotype of Calumza biocellata, Zanzibar. AMS I. 19472-086, 2(14-19) , Yonge Reef, Great Barrier Reef, Australia, 7-15 m. I. 19480- 026, 2(19-22) , Yonge Reef, Great Barrier Reef, Australia, 20-30 m.
Calumza profunda sp. nov. Figs 2, 3 and 4 Diagnosis: Mouth enlarged, reaching to below middle of eye or slightly beyond. Gill opening broad, extending forward to below middle to posterior end of pupil. Elongate rakers on outer face of first gill arch 13-14. Pectoral rays 14-15. Lateral scale count 24- 25. Transverse scale count (TRB) 8. Second spine of first dorsal fin the longest and slightly prolonged. Seven dark brown transverse bands on body. Caudal fin without
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
20 SPECIES OF CALUMIA
enlarged black spots. Anterior tip of tongue bilobed. Pelvic rays unbranched. Outer row of jaw teeth not enlarged.
Description: Based on holotype and two paratypes. Counts of holotype indicated with an asterisk. Measurements of types given in Table 1. First dorsal rays VI (3) *. Second dorsal rays I, 6 (1); I, 7 (2)*. Anal rays I, 7 (3)*. Pectoral rays 14 (1), 15 (2)*. Lower rakers on outer face of first arch 13 (2)*, 14 (1). Segmented caudal rays 14 (1), 15 (2)*. Lateral scale count 24 (1), 25 (2)*. Transverse scale count (TRB) 8 (3) *. Predorsal scales 8 (3) *.
A small species maturing at about 15 to 18 mm SL. Body compressed posteriorly, more rounded anteriorly. Head large, 36 to 39% of SL, distinctly depressed. Eyes lateral, top of eye forming top of head profile, about 4 in head length. Jaws terminal, slightly oblique; end of jaws below posterior half of pupil or below midpupil. Tip of lower jaw protrudes slightly before tip of upper jaw. Anterior nostril an elongate tube about equal to pupil diameter, positioned just behind upper lip. Posterior nostril a raised rim midway between anterior nostril and front of eye. Snout about equal to eye, with slight protuberance on dorsal profile before eye.
Dorsal and anal fins short based, shorter than caudal peduncle length. First dorsal fin slightly elongate with a pointed distal margin, formed by the elongate second dorsal spine. Dorsal and anal rays elevated becoming progressively longer posteriorly, giving fins pointed posterior margins; tips of fins reach to base of caudal rays. Pectoral rays slender, unbranched, middle rays longest reaching to above anterior part of anal fin. Pelvic fins separate, composed of one spine and five unbranched rays; first segmented ray short, rays becoming progressively longer, with fourth ray extending to below middle to end of anal base; fifth segmented ray short, about equal in length to first ray. Caudal fin short, oval in shape, shorter than head length. Gill opening broad, with branchiostegal membranes attaching to middle of isthmus below middle of eye. Body scales ctenoid, but cycloid on head, breast, belly and pectoral base. Cheek and opercles covered with large scales. Scales on top of head extend forward to above the end of the eyes and a single enlarged scale between eyes. Both jaws with a band of irregularly spaced small curved pointed teeth, and an innermost row of larger, straight, backward-pointed teeth; bands of teeth extend full lengths of jaws, but become narrower posteriorly. No vomerine or palatine teeth. No
TABLE 1
Measurements of types of Calumza profunda and recently collected material of C. godeffroy: in millimetres
Measurement C. profunda C. godeffroyz Holotype Paratype Paratype AMS I 19472-086 BPBM AMS I AMS I 21158 17477-026 _20156-001 Sex °} 2 g fo) } Standard length 18 16.5 17 14 19 Head length 7.0 6.0 6.7 4.9 6.8 Head depth at preopercular margin 3.8 3.0 3.3 3.0 2.6 Head width at preopercular margin 4.3 3.8 4.0 2.9 2.9 Upper jaw length 3.0 2.8 2.8 1.6 1.9 Eye length 1.8 1.6 1.6 1.5 1.7 Body depth at analorigin 4.0 3.4 3.5 3.6 4.5 Caudal length = 4.8 4.8 3.8 =
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
H.K. LARSON AND D.F. HOESE 21
Fig. 2. Holotype of C. profunda. Drawing by H. K. Larson (caudal fin reconstructed).
NS em
vg @ REPP,. @eov)sesv® 9g
Fig. 4. Top (right) and ventral (left) views of head of C. profunda showing arrangement of sensory papillae.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
22, SPECIES OF CALUMIA
open lateral line head pores. Sensory papillae sparse, obscured by scales on cheek, but distinct below eye, on top of snout and on lower surface of head (Figs 3, 4).
Colouration in alcohol: Head and body light brown. Centres of scales dark brown. Seven broad bands on body; first behind pectoral base; second under first dorsal fin; third and fourth under second dorsal fin; fifth and sixth on caudal peduncle; seventh at base of caudal. Head with faint brown bars radiating from eye ventrally and posteriorly. Snout, lips and underside of head dark brown. A dark brown blotch on each side of nape above upper end of operculum in front of pectoral base. Pectorals and pelvics with scattered melanophores. A dark brown spot at base of each pelvic fin. Caudal fin uniformly dusky, darker above and below base, where seventh body band extends onto fin. First dorsal fin with broad dark basal band, rest of fin clear. Second dorsal and anal fins dark brown basally, with body bands extending onto bases of fins; upper third of fins clear.
Colouration of freshly collected holotype: Head and body chocolate brown. Iris golden yellow. Nostrils white. Head with three orange stripes radiating posteriorly from eye. Posterior third of operculum and branchiostegal membranes orange. Scattered orange mottling on sides and top of head. Body bands chocolate brown, interspaces light brown anteriorly, white posteriorly. Scale centres orange in dark body bands only, forming distinct rows of small orange spots, about half of pupil diameter in size. Base of first dorsal bluish with scattered small brown and orange spots; middle of fin with a broad bright yellow stripe; tip of fin bluish. Second dorsal and anal as for first dorsal, except basal two thirds of fins with large orange spots and yellow stripe above middle of fins. Caudal yellowish above and below, middle of fin clear, with tiny white speckles; two or three small white spots along upper and lower margins near base of fin. Pectoral and pelvic fins clear to whitish. Base of pelvic fin with a black spot.
Etymology — from Latin, profunda — of the depths, alluding to its being found about relatively deep coral reefs.
Material Examined — Holotype: BPBM 21158, an 18mm female from 38 m depth Alite Reef, near Malaita, Solomon Islands; J. Randall and B. Goldman, 25 July 1973. Paratypes: AMS I.20156-001, a 17mm female, taken with holotype. I.17477-026, a 16.5mm female from 55m depth, Bogacio Island, Espiritu Santo, New Hebrides; G. Allen, W. Stark and D. Popper, 28 June 1973.
ACKNOWLEDGEMENTS
We would like to thank J. E. Randall for supplying some of the type material and a colour photo of the holotype. We would also like to thank M. M. Smith (RUSI) and A. Wheeler (BMNH) for making types of described species available. J. R. Paxton kindly reviewed the manuscript.
References
ALLEN, G. R., 1979. — The fishes of Christmas Island, Indian Ocean. Aust. Nat. Parks Wildlife Spec. Publ. 2: 1-81.
GUNTHER, A., 1877. — Andrew Garrett’s Fische der Sudsee, Part 6. J. Mus. Godeffroy 4 (13) : 169-216.
Husss, C. L., and Lacier, K. F., 1958. — Fishes of the Great Lakes region. Bloomfield Hills, Michigan: Cranbrook Institute of Science. 213 pp.
SCHULTZ, L. P., 1966. — Fishes of the Marshall and Marianas Islands. Volume 3. Bull. U.S. Natn. Mus. 202: 1-165.
SMITH, J. L. B., 1958. — The fishes of the family Eleotridae in the western Indian Ocean. Ichth. Bull. Rhodes Univ. 1: 137-163.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
The Eelgrass Zostera capricorn in lawarra Lake, New South Wales
M. MCD. HARRIS, R. J. KING and J. ELLIS
Harris, M. McD., Kine, R. J., & Eis, J. The eelgrass Zostera capricornt in Illawarra Lake, New South Wales. Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980:23-33.
Zostera capricorni Aschers. is the dominant angiosperm in Illawarra Lake where it occurs over a wide range of substrate types (14-98% sand, 0.5-12% organic carbon) and salinity (3 % — normal seawater). Distribution and abundance are related to light availability. The growth cycle shows a maximum in summer, with a winter minimum following shedding of the previous season’s growth. Flowering is extensive but even though seedlings have been observed, propagation appears to be almost entirely vegetative. Zostera, complete with substrate, has been successfully transplanted.
M. McD. Harris, Wollongong Institute of Education, Wollongong, Australia 2500; R. J. King, School of Botany, University of New South Wales, Kensington, Australia 2033; and J. Ellis, Department of Chemistry, University of Wollongong, Wollongong, Australia 2500; manuscript received 22 November 1978, accepted in revised form 21 March 1979.
INTRODUCTION
Illawarra Lake is a coastal saline lagoon which straddles the boundary of the city of Wollongong (34°30'S, 150°50’E) in the north, and the Shellharbour municipality in the south, Fig. 1. The lake has evolved from a broad bay by the formation of a baymouth sand bar (Thom, 1974).
Lakes formed in this way tend to be broad, shallow, exposed and turbulent expanses of water with an extensive sandy zone supporting benthic macrophytes. Illawarra Lake is no exception. It has a maximum length of approximately 9.5 km and a width of 5.5 km with an approximate area of 33 km’. The lake is shallow; an estimated 25% is less than 1.2 m deep and the maximum depth is only 3.7 m (Roy and Peat, 1973). The lake is oriented N-S parallel with the coast and is thus exposed to strong southerly and south westerly winds; lake water can be turbulent with wave heights approaching 0.5 m (Eliot et al., 1976). This turbulence coupled with the shallow nature of the lake causes considerable turbidity and is probably a significant factor in maintaining the concentration of dissolved oxygen (Kanamori, 1976).
At present the lake enters the ocean near the southern end of the sand barrier where it is partially protected by Windang Island. This rocky prominence, which is usually land-tied by a tombola, dissipates wave energy and thus suppresses the rate of infilling of the entrance channel. The present channel from the lake to the ocean changes continually in position, width and depth depending on factors such as rainfall, wind and wave action. During 1972-1977 the entrance channel has been about 2.5 km long, winding and varying in depth up to 2.5 m. The main channel is seldom more than 100 m wide. It is restricted by a sand bar at the Windang bridge and occasionally the bar has completely choked the entrance. In late 1971 a blocking bar formed and this was subsequently cleared with earthmoving equipment. As a consequence of the restricted access to the sea very little tidal influence extends into the lake even at spring tides. Eliot et al. (1976) report a tidal rise of up to 0.1 m (cf. 2.0 m on the nearby ocean coast) on the western side of the lake at the Tallawarra power station. Ellis et al. (1977) estimate a tidal volume of 1% of the lake volume.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
24 ZOSTERA CAPRICORNI IN ILLAWARRA LAKE
SL Ck, is} \
Mullet Ck.
Brooks Ck.
Kanahooka Pt.
Koonawarra Bay
Tallawarra Pt.
Windang Is.
Karoo Bay
Fig. 1. Map of Illawarra Lake with geographical names
THE FLORA OF ILLAWARRA LAKE
Zostera capricorni Aschers. is the only seagrass recorded for Illawarra Lake and its distribution and abundance are the main subject of this paper. The range of morphology exhibited by Z. capricorni in Illawarra Lake is considerably greater than that given by Den Hartog (1970). Ruppza sp. is the only other angiosperm abundant in the benthic flora: it occupies approximately 30% of the eastern weed beds as a dense meadow in water 40-60 cm deep, Fig. 2. In shallow areas growth is reduced to sparse clumps. The change to Zostera with deeper water is abrupt. Isolated patches of Ruppia occur throughout the lake and in Hooka Creek, over a salinity range of 3-329. At all localities Ruppza grows inshore of Zostera, the reverse of the situation outlined by Higginson (1965) in the Tuggerah Lake system. Higginson suggested that Ruppia tended to favour clayey sediments, cf. Zostera on sandy sediments, but this conclusion was not borne out in this study. Wood (1959a) and Higginson (1965) both concluded that Ruppza is intolerant of strong currents and occurs mainly in sheltered bays. In this case high turbulence in Illawarra Lake may be the limiting factor for Ruppia which is then confined to the sheltered localities inshore of Zostera.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
M.M. HARRIS, R.J. KING AND J. ELLIS 25
Eastern Weed Beds
a |Windang
Cudgeree
Entrance Channel
Haywards Bay
=: Zostera a Ruppia ® Lamprothamnio 4 Gracilaria \
Fig. 2. Distribution of benthic plants in Illawarra Lake
Posidonza australis Hooker is not recorded in Illawarra Lake though the reasons for its absence remain obscure. Poszdonza is reported to be intolerant of strong currents and turbulence (Wood 1959a, b; Den Hartog, 1970) and also appears to be intolerant of high turbidity. High turbidity may prevent growth of Pos¢donza in the body of the lake but could not account for its absence in the entrance channel area. Turbulence and current velocities in the channel area do not approach the rip conditions observed in the entrance to Macquarie Lake where Wood (1959a) reports growth of Poszdonza.
Halophila ovalis (R. Brown) Hooker was also surprisingly absent from Illawarra Lake. It occurs along the N.S.W. coast on a wide range of sediment types encompassing those found in Illawarra Lake, and over a wide range of conditions of salinity and turbidity. Despite its apparent wide ecological tolerance Halophila has not been observed in Illawarra Lake.
Macroalgae were common in the Lake, particularly as epiphytes on Zostera and Ruppia. The most conspicuous were Polysiphonia sp., Enteromorpha intestinalis (L.) Link and Cladophora sp.; the latter species generally occurring in spring and early
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
26 ZOSTERA CAPRICORNI IN ILLAWARRA LAKE
summer. The Enteromorpha and Cladophora form extensive floating mats in protected bays and in the inshore parts of the eastern weed beds. The benthic alga Gracilaria verrucosa (Huds.) Pap. occurred throughout the year and the broad distribution is shown in Fig. 2. In the southern bays, Koona and Haywards Bays, it was the dominant species occupying the silty sediments in the central portion of the bays. At least in initial stages plants are attached to the sediment. Lamprothamnion was variable in occurrence but mainly associated with Ruppza inshore of Zostera in the eastern weed beds, Fig. 2.
METHODS
The data and observations presented in this paper were gathered over the period 1971-1976. Complete details can be obtained from Harris (1977). The data include: observations on the distribution and abundance of Zostera capricorni and other components of the benthic flora including epiphytes; details of the relationship between Zostera distribution, biomass and environmental data; long term observations on growth and flowering in Zostera.
In the analysis of sediments and water quality the following techniques were used :
Particle size analyses were conducted according to Folk (1968) .
Organic carbon was estimated from loss on ignition at 550°C as described by Dean (1974).
Total phosphorus was determined colorimetrically using the single solution method detailed in Major et al. (1972).
pH was measured using a specific ion meter and pH combination electrode (Orzon Model 407A and electrode no. 91-02). Sediment pH is based on interstitial water extracted from the sediment by pressure.
Eh was measured using an Eh combination electrode (Orion no. 96-78) standardized using Zo Bel solutions as described by Whitfield (1971). Sediment Eh was made on carefully mixed samples.
Salinity was measured as chlorinity using a chloride electrode (Orion no. 96-17). Temperature was measured using a standard 50°C mercury thermometer.
Turbidity measurements were initially made with a secchi disc but later using a turbidimeter (Hach Model 2100A).
Wherever possible data from other sources, particularly Ellis and co-workers, were used.
RESULTS AND DISCUSSION I Distribution of Zostera in Illawarra Lake in relation to environmental factors
The major development of Zostera capricorni occurred in the eastern portion of the lake, and particularly off the Windang Peninsula and west of Bevans Island, Fig. 2. A narrow fringing zone seldom more than 20 m wide occurred around most of the bays in the western portion, except where Graczlarza was dominant.
(A) Sediment Factors. The composition of sediments supporting Zostera is given in Table 1. Extreme ranges were: particle size, sand 52 to 98%; organic carbon 0.5 to 8.5%; total phosphorus 35 to 120 wg.g™'. Even greater extremes in values for particle size and organic carbon relate to the single site in Hooka Creek: 14% sand and 12% organic carbon. pH and Eh measurements were restricted to the eastern sandy portion of the Lake. Sediment pH ranged from 7.4 to 8.0 and Eh from +10 to —185 mV. This range of sediments, together with the lack of relationship between the distribution of sediment types in Illawarra Lake and the distribution of Zostera suggest that Zostera is not limited by sediment type.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
M.M. HARRIS, R.J. KING AND J. ELLIS 27 TABLE 1
Composition of sediments supporting Zostera in Illawarra Lake Mean values: range given in parentheses
Location Number of samples Sand % Organic carbon % _Total phosphorus yg.17? Griffins Bay 12 90 (77-97) 2.3 (0.8-3.5) 82 (35-105) Eastern Weedbeds
(off Windang Pens.) 8 96 (95-97) 1.5 (0.5-2.7) 61 (39-111) Bevans Island 18 92 (88-97) 1.8 (0.8-4.3) 72 (53-120) Southern Bays 5 82 (52-97) 3.3 (1.3-8.5) _
Western Weedbeds 5 84 (58-92) 353) (Zhe 522) -
Hooka Creek 1 14 12.0 —
Where the reduction of Zostera beds occurred in Koona Bay and Koong Burry Bay over the period 1972 to 1977 it appeared to be related to rapid accretion of sediments. In other places in the lake (e.g. the northern side of the entrance channel east of the Windang Bridge, on the delta at the lake end of the entrance channel, in the shallows to the north of Cudgeree Island and the delta of the Griffins Bay tank trap) burial and subsequent recovery has been a frequent occurrence. (B) Water Factors. Water quality in Illawarra Lake was highly variable. This is partially due to the fluctuating level of the lake: records of the N.S.W. Electricity Commission (1971) show an average variation in lake level of about 60cm in the period 1966-70.
Salinity. Ellis et al. (1977) showed that mean salinity in the Lake is controlled primarily by rainfall, and varied from 12.8 to 31.3°%, during a two year monitoring period. There was essentially no salinity stratification, with vertical mixing by wave action completed within 2-4 weeks. There was no significant east-west salinity gradient even though the major creeks are on the western side of the lake.
Zostera grew and even flowered over a wide range of salinity, from approximately 3%» in Hooka Creek to approximately 35 9%, in the entrance channel. This is outside the range experienced in the lake and it was concluded that salinity did not limit Zostera distribution.
Temperature. Ellis and Kanamori (1977) record a mean water temperature for Illawarra Lake in the range 11.6 to 25.6°C. In the Tallawarra Power Station outlet channel, water temperatures at times exceeded 35°C yet Zostera persisted at this site. It is concluded that water temperature was not a limiting factor since the differences in temperature between various sites on the lake was always slight, usually less than 3°C, and new growth was observed throughout the year.
Phosphorus. The nutrient status of the lake water, with regard to phosphates was generally beyond the level of enrichment needed to promote algal blooms (Anon., 1975; Wetzel, 1975). The observed total phosphorus range was 4 to 145 wg P.1” (Kanamori, 1976). Because the lake was turbulent and the phosphate level of the sediment high, localized deficiencies in water phosphorus concentrations would be quickly restored by water circulation and by disturbance of the sediment. It is thus unlikely that the distribution would be limited by phosphorus deficiency.
Some very high concentrations of total phosphorus, up to 3,600 yg.1™' were observed in small creeks draining from non-sewered urban areas. Associated with this was a high E. cold count indicating that much of the nutrient inflow to Illawarra Lake is derived from domestic effluents and sewage. In areas adjacent to some streams, and in some bays, notably near Albion Creek in Koona Bay, in Kully Bay and Joes Bay
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
28 ZOSTERA CAPRICORNI IN ILLAWARRA LAKE
(Griffin Bay), Why Juck Bay, and in the Back Channel, Zostera grew sparsely or not at all. In these areas the green alga Enteromorpha grew in large floating masses shading the Zostera. In this way excessive phosphorus levels may have been indirectly responsible for restricting the distribution of Zostera.
Nitrate levels in the Lake have a mean value of 2 wg.1-* (Kanamori, 1976) which is lower than values reported by Higginson (1971) for Tuggerah Lakes or Spencer (1959) for Macquarie Lake.
pH and Eh. The observed ranges of pH and Eh for lake water were narrow; pH 8 to 8.4 and Eh +310 to +350 mV. State Pollution Control Commission figures (unpublished) give a pH range of 7.4 to 8.9. The narrow range and lack of systematic variation between different areas of the lake suggest that it is improbable that either of these factors would limit Zostera within the lake.
Turbidity. Light availability has often been demonstrated as the factor setting the maximum depth to which seagrasses will grow (Backman and Barilotti, 1976). In Illawarra Lake Zostera occurred to a depth of 2 m in the entrance channel which was twice daily filled with low turbidity sea water. Throughout most of the eastern weed beds the outer limit was usually found at 1.5 to 1.8 m even though the sandy substrate continued beyond 2m in depth. In Griffins Bay and off the mouth of Mullett Creek, areas of fine sediment and high turbidity, the limit was inside the 1 m contour.
Turbidity values and the depth limit of Zostera growth for near Bevans Island, the eastern weed beds, and one site in Griffins Bay are given in Table 2. Values are given for turbidity of water on the outer edge of the seagrass beds and also within the seagrass bed. Turbidity measure can often change rapidly: in the narrow fringing seagrass beds on the western side of the lake, where sediments are generally finer, turbidity could change from 2 to 40 NTU (nephelometric turbidity units) within 10 min. of a strong southerly change acting upon the lake. In the highly turbid areas of Koong Burry Bay, Haywards Bay and Koona Bay, Zostera is dominated by Gracilaria, in Griffins Bay Zostera and Gracilaria are co-dominant and in the eastern weed beds Gracilarza is only of minor importance.
Water Movement. Zostera is reported to grow in regions with considerable water movement (Higginson, 1965). In Illawarra Lake it appeared to tolerate currents of several knots but to be less tolerant of strong wave action. Extensive colonies occurred on the margins of the entrance channel where they experienced strong tidal flow, and colonies in Mullet and Hooka creeks were not obviously disturbed by flood flows.
The role of wave action is difficult to assess. On exposed peninsulas like Wollingurry Point, Tallawarra Point, Kanahooka Point and Wollamai Point, or in
TABLE 2
Turbidity measure at various sites in Nephelometric Turbidity Units
SITE
Near Bevans Island Eastern Weed Beds Griffins Bay
Depth of weed growth 1.8m 1.8m 1.6m 1.0m
Turbidity measure atmargin in weed bed
10.11.75 2.0 0.5 1.5 4.5 1.75 1.0 4.5 9.5 8. 1.76 1.5 3.5 1.0 2.0 2.0 2.0 3.0 3.5 17. 1.76 1.5 15 1.5 1.5 2.0 2.0 2.5 1.5 29. 1.76 1.5 2.75 Wey 0) Neds. Boe 2.0 3.0 8. 2.76 3.0 1.25 1.5 2.5 2.0 1.5 2.757 9350. 14. 2.76 3.5 3.5 3.0 4.75 3.0 3.0 4.5 3.0 Mean Zee 2.0 1.6 2.9 Zell 2.0 3.2 3.9
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
M.M. HARRIS, R. J. KING AND J. ELLIS 29
open bays like Tuggerah, Yallah and Moureendah Bays, strong wave action periodically deposited several centimetres of sediment; then later removed it exposing the underlying rock. Zostera was unable to produce more than minor colonies in these areas.
During 1975-76 the Zostera beds in the northern side of the training wall in Yallah Bay degenerated greatly. This occurred during a period of high wave action generated by persistent, strong easterly winds and considerable sediment disturbance was evident. Similar observations were made in the beds south of Wollingurry Creek.
Of the environmental factors examined water depth and turbidity appear to be the most important factors limiting the lower distribution of Zostera in the lake. There is no evidence that sediment type plays any major role as has been suggested in the case of submerged aquatic angiosperms in the Tuggerah Lakes system (Higginson, 1965). The minimum water depth limit for Zostera in Mlawarra Lake varied widely. In Griffins Bay, along the eastern weed beds and west of Bevans Island Zostera was sparse in water less than 40 cm deep. These are areas of intensive feeding by ducks and swans. The ducks have been observed feeding on the shoots of Zostera and the swans on the rhizomes of both Zostera and Ruppia. Im areas such as the Back Channel, swans were seldom observed and here Zostera grew well in 15 - 20 cm of water. In September 1976, when lake levels were low Zostera beds in Koonawarra Bay were exposed. During this period several hundred black ducks (Anas superciliosa) congregated in this area and over the next 3 days cropped the exposed Zostera and Ruppia to within 1 cm of the sediment surface.
II Zostera biomass in relation to environmental factors
Analysis of the transect biomass data with environmental data showed no overall correlation between Zostera biomass and the factors particle size, sediment pH or Eh, ‘water pH or Eh, water temperature, and water or sediment total phosphorus. Despite the greater nutrient content of sediments in Griffins Bay, compared with those near Bevans Island, Zostera biomass at the latter location was significantly higher (1.5 - 3 times) than in Griffins Bay. There was however a distinct relationship between Zostera biomass and water depth, and hence light availability, Fig. 3a. Fig. 3b shows the growth of Zostera expressed as shoot length at the same sites. A similar relationship holds for total plant biomass and water depth. The differences between Bevans Island and Griffins Bay data are consistent with the notion that depth limits of Zostera at a given locality are a function of turbidity. The reduced biomass in water less than 40 cm is presumed to relate to waterfowl grazing pressure.
III Vegetative growth of Zostera capricorn
The seasonal growth cycle data for Zostera at selected localities during 1972-73 are shown in Fig. 4. The general pattern is similar with maximum leaf length occurring during summer and the minimum in winter. The onset of the new growth cycle occurred in August - September after shedding of the previous season’s growth. Wood (1959a) reported that leaf loss follows flowering and in autumn Zostera flats may seem completely bare of leaves. In this study full leaf development was sometimes maintained until early spring by which time new leaf growth had commenced.
In 1973 rapid degeneration of Zostera beds occurred during February and March coincident with heavy rainfall. Similar declines in the standing crop followed heavy late summer and autumn rains in 1974 and 1976. In 1975 when rainfall was 40% less than average little decline occurred until the June - July floods. These changes were most marked in waters shallower than 0.6 m. This relationship between flood rains
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
30 ZOSTERA CAPRICORNI IN ILLAWARRA LAKE
200
A Griffins Bay
@® Bevans Island
Zostera biomass
40 60 80 100 120
Water Depth (cm) Fig. 3a. Zostera biomass (mean values) in relation to water depth — Bevans Island and Griffins Bay
Modal Zostera shoot length (cm)
40 60 80 100 120
Water Depth (cm)
Fig. 3b. Zosterc shoot length (mean values) in relation to water depth — Bevans Island and Griffms Bay, January 1976
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
M. M. HARRIS, R.J. KING AND J. ELLIS 31 100
EASTERN WEED BEDS
80
60
40
20
100 NEAR BEVANS ISLAND
Zostera shoot length (cm)
100
HAYWARDS BAY
80
60 j_ 40
20
Jul Aug Sep Oct Nov Dec Jan 1973
Fig. 4. Seasonal variation in shoot length of Zostera capricorni showing modal shoot length and range.
and leaf shedding was most marked in the eastern weed beds while colonies in
sheltered localities such as Haywards and Koonawarra Bays or along the southern
shore were barely affected. When the Zostera beds have their normal water cover,
wave action is suppressed at the outer margin; however, during flooding the increased
water depth allows for vigorous wave action throughout the beds, leading to extensive _ shedding of leaves. Associated lower salinities and increased turbidity may predispose
Zostera towards leaf shedding but in the absence of mechanical agents it did not
appear to arise from these causes alone.
IV Flowering and Seed Production
Flowering of Zostera in Illawarra Lake is variable both temporally and spatially. Immature flowering shoots were first observed in September and flowering sometimes extended through to August of the following year, with seed production from October to August. Generally peak flowering occurred in summer. Table 3 summarizes data for the season 1975-76. In that season flowering was abruptly terminated with
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
32 ZOSTERA CAPRICORNI IN ILLAWARRA LAKE TABLE 3
Percentage of fertile shoots at three locations in summer 1975-76 based on 500 cm? composite sample
Number of shoots % fertile shoots Number of seeds
examined
Site October 1975 November December January 1976 February Near BevansIsland — nil — 992 4.1 57 650 10.6 89 579 13.9 112 406 2.2 7 Eastern Weed Beds 136 6.6 — 1295 7.5 34 679 12.2 61 516 17.2 83 250 1.2 2
Griffins Bay — nl — 257 22.2 186 397 12.1 172 405 13.6 101 — nil —
shedding of flowering shoots after heavy January rains. The frequency of flowering shoots showed wide variation between years: in 1972-73 the frequency reached 34% near Cudgeree Island and in Haywards Bay.
The majority of flowers did not produce seed. The observed maximum of flowers producing seed was 10.9% in 1976 compared to less than 2.6% in 1973.
Germinating seeds were found in sieved sediment samples on only one occasion: near Bevans Island in July 1975. There was no evidence that Zostera was colonizing bare areas by seedlings. In dredged areas any subsequent recolonization occurred by rhizome invasion from adjacent areas. As found by Wood (1964), such recolonization is slow. Preliminary experiments with transplanting Zostera into the back channel near the Tallawarra Power Station and in Griffins Bay were successful when the plants were transplanted complete with relatively undisturbed sediments: washed rhizomes did not recolonize. These small transplants (size range 100 - 200 cm”) were made in June - October 1975 and 3 years later those near the Tallawarra Power Station had expanded to cover some 10 times the original area. This expansion has taken place by marginal growth of the clump into formerly uncolonized sediments. Growth in Griffins Bay has been less pronounced though clumps are still growing.
CONCLUSION
Within Illawarra Lake Zostera capricornt showed a marked ecological tolerance. Within the range found in the lake the distribution and abundance of Zostera did not appear to be controlled by substrate factors (particle size, total phosphorus, pH, Eh) or water quality factors (salinity, total phosphorus. pH, Eh). Light availability as influenced by water depth and turbidity appeared to control the lowest depth to which the plant grows. Grazing by swans and ducks may have been significant in setting the upper limit.
Zostera showed a seasonal cycle with shedding of the previous season’s growth promoted by wave action. Flowering was extensive, though patchy and seed production was observed over a lengthy period, October to August of the following year. Although seedlings were observed in the field there was no evidence to suggest that seedlings play a major role in propagation. Transplant experiments, /using Zostera clumps in relatively undisturbed sediments, have been successful and clumps have been expanded by growth into previously uncolonized sediments.
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
M. M. HARRIS, R.J. KING AND J. ELLIS 33
ACKNOWLEDGEMENTS
We wish to acknowledge grants from the N.S.W. Electricity Commission, and the Department of Urban and Regional Development, under a National Estate Grant provided to the Wollongong City Council for the Illawarra Lake Environmental Assessment Project.
References
Anon., 1975. — Re-using sewage water. Ecos 3: 14-18.
BACKMAN, T. W., and BariLotTI, D. C., 1976. — Irradiance reduction: effects on standing crops of the eelgrass Zostera marzna in a coastal lagoon. Mar. Bzol. 34: 33-40.
DEAN, W. E., Jr., 1974. — Determination of carbonate and organic matter in calcareous sediments and sedimentary rocks by loss on ignition: comparison with other methods. J. Sedim. Petrol. 44: 242- 248.
DEN HartToc, C., 1970. — The Sea-Grasses of the World. Amsterdam: North Holland Publ. Co.
ELECTRICITY COMMISSION OF N.S.W., 1971. — Tallawarra area climatic and limnological data. (Unpublished) .
Exiot, I. G., Younc, R. W., and Crark, D. J., 1976. — Part 4 — Lake hydrology. In Illawarra Lake. Wollongong: Wollongong City Council.
Exuis, J., and KANAMORI, S., 1977. — Water pollution studies on Lake Illawarra. II. Thermal behaviour. Aust. J. Mar. Freshw. Res. 28: 479-484.
——, Kanamori, S., and Lairp, P. G., 1977. — Water pollution studies on Lake Illawarra. I. Salinity variation and estimation of residence time. Aust. ]. Mar. Freshw. Res. 28: 467-477.
Fok, R. L., 1968. — The Petrology of Sedimentary Rocks. Austin, Texas: Hemphills.
Harris, M. M., 1977. — Ecological studies on Illawarra Lake with special reference to Zostera capricorni Ascherson. Kensington: University of N.S.W., M.Sc. thesis, unpubl.
Hiccinson, F. R., 1965. — The distribution of submerged aquatic angiosperms in the Tuggerah Lakes system. Proc. Linn. Soc. N.S.W. 90: 328-334.
——, 1971. — Ecological effects of pollution in Tuggerah Lakes. Proc. Ecol. Soc. Aust. 5: 143-152.
KANAmorI, S., 1976. Water pollution studies on Lake Illawarra. Wollongong: University of Wollongong, Ph.D. thesis, unpubl.
Major, G. A., Dat Pont, G., Kye, J., and NEWELL, B., 1972. — Laboratory Techniques in Marine Chemistry. CSIRO Div. Fish. and Oceanogr. Rep. 51.
Roy, P. S., and Peat, C., 1973. — Bathymetry and bottom sediments of Lake Illawarra. Geol. Surv. N.S.W., Report No. GS 1973/295.
SPENCER, R. S., 1959. — Some aspects of the ecology of Lake Macquarie N.S.W.., with regard to an alleged depletion of fish. II. Hydrology. Aust. J. Mar. Freshw. Res. 10: 279-296.
Tuom, B. G., 1974. — Coastal erosion in eastern Australia. Search 5: 5.
WETZEL R. G., 1975. — Limnology. Philadelphia: Saunders.
WHITFIELD, M., 1971. — Ion selective electrodes for the analysis of natural waters. AMSA Handbook No. 2. Sydney: Aust. Mar. Sci. Assocn.
Woop, E. J. F., 1959 (a). — Some east Australian sea-grass communities. Proc. Linn. Soc. N.S.W. 84: 218- 226.
——, 1959(b). — Some aspects of the ecology of Lake Macquarie, N.S.W., with regard to an alleged depletion of fish. VI. Plant communities and their significance. Aust. J]. Mar. Freshw. Res. 10: 322- 340.
——., 1964. — Studies in microbial ecology of the Australian region. Nova Hedwigza 8: 5-54, 453-568.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
A Key to Estuarine Polychaetes in New South Wales
PAT HUTCHINGS and SEBASTIAN RAINER
Hutcuincs, P., & RAINER, S. A key to estuarine polychaetes in New South Wales. Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980:35-48. A key to 184 species of estuarine polychaetes in New South Wales is given together with a reference to an adequate description of each named species.
Pat Hutchings, Dept of Marine Invertebrates, Australian Museum, P.O. Box A285, Sydney South, Australia 2000, and Sebastian Rainer, CSIRO Division of Fisheries and Oceanography, Cronulla, Australia 2230; manuscript received 21 February 1979, accepted in revised form 18 July 1979.
INTRODUCTION
The estuarine polychaete fauna of New South Wales includes many undescribed species and new records. Sheltered bays will share many species in common with estuarine areas. At present we know of 184 species occurring in N.S.W. estuaries of which 125 have been described. Many other species remain to be described, some perhaps belonging to families not included in this key, for example Chrysopetalidae and Amphinomidae. Many of the descriptions were published early this century in journals with limited circulation and often not in English. No key exists to enable most of these species to be identified or differentiated from species already described and this key is intended to remedy this situation.
The key is based on material collected during surveys of Careel Bay (Pittwater) and Gunnamatta Bay (Port Hacking) by the authors, from cores collected from Posidonia beds along the coast of New South Wales (PH and others) and other material from the Australian Museum collections. It is likely that many more species will be found in estuarine situations in New South Wales, but the key should provide a reasonable guide to at least the larger species. As it is possible that closely related species not in the key could be confused with keyed species, identifications made with the key should be checked against the literature. The number in brackets accompanying each named species indicates where an adequate description of that species may be found, using the numerical order of the references cited. Species keyed only to genus may represent undescribed species, new records that we have not been able to confirm or may refer to incomplete material that did not permit positive identification. An indication of other recorded species from Australia that may occur in N.S.W. estuaries is given by Day and Hutchings (1979).
The key is based on Day (1967) with some modifications and additions using the taxonomy of Fauchald (1977). Reference should be made to Day (1967), for diagrams indicating the important features of individual families and for a glossary of the terms used.
KEY TO ESTUARINE POLYCHAETES IN N.S.W. (modified after Day, 1967)
1. —Most of the following characters: prostomium with
sensory appendages; pharynx armed with jaws or
teeth; parapodia well developed, compound setae OLLENIPTESENE Wiese ewe Ackles ee sheaers leh ey Mine mnewsieanac) alls Polychaeta Errantia 2
—Most of the following characters: prostomium usually
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
36 ESTUARINE POLYCHAETES
lacking sensory appendages, often fused to peristome, which may bear grooved palps, buccal cirri, stout setae or branchial crown; compound setae rarely
PRESENT Ae OI aN E NED. MGs ay ara ame ae cM Polychaeta Sedentaria 68 Polychaeta Errantia 2h —Elytra present on many segments ............... .e eee ee eee eee eee 3 == Fiytravalbsem tie iitascasieel syealtoviot onan cisstcuianeneh cies ee lene, cri amystcch cor ente tutes) cite trae veti eS RAL aR a ge US 12 3. —Compound setae absent; elytra and dorsal cirri alternate fairly regularly...................... F. Polynoidae 4
—Compound setae present; elytra on alternate segments anteriorly, from about setiger 25 onwards Onl allisegments ie 253k cece ain a ea OU Gr Stn yi F. Sigalionidae 11
4. —Lateral antennae terminal, arising at level of median antenna, 15 pairsofelytra .................... Parahalosydna chrysostichtus (13) —Lateral antennae arising below level of median antenna’) 5-16) pairs of elytraiy hy ala’ ie ins Weiter ils rales peteere trae netnrn et alvel etesy Mi lurealbal Mea 5
5. —Lateral antennae subterminal;. ventral lamellae PLESEM lich saeco ossis tar iyrea sts ce tue Sle: obs aa mucus scuanay eieal Shoat tes Paralepidonotus ampulliferus (13) —Lateral antennae ventral; ventral lamellae absent... .....................000% 6
6. —Neurosetae with blades tapering to fine tips, UMI GEN tates sas te won itn aon aaron ane ch alade Antinoe sp.
—Neurosetae with blades ending in stout tips, uni- or bidentate {occ woe nana nant me cee MeN dhe fal ihe ea DI anda oh gts SMR Cony Ne anvL Santa 7
Pp —One or more of the basal serrations on both notosetae and neurosetae enlarged to form spinous pockets; prostomium without frontal peaks .............. Scalisetosus sp. — Basal serrations on setae not enlarged, prostomium with frontal peaks.....................00008, Harmothoe 8
8. —Elytra ornamented with multifid tubercles ........ Harmothoe sp. —Elytra ornamented with simple tubercles.......... 2.2... eee eee eee 9
9. —Elytral tubercles with flat-topped apices .......... Harmothoe sp. —Elytralltuberclesiconical: 2/95 foe eee n poenalsha sh Yess aieesiciel euenaamue nl areasieusiinne ests 10
10. —Elytral tubercles numerous.................... Harmothoe sp. —Elytral tubercles short, conical, few, sometimes absent, elytra, margins with elongate papillae...... H. praeclara (13)
11. — Median antenna with basal lappetson ceratophore.. Sthenelazs sp. —Median antenna absent or papilliform and lacking CET atOphore ss site doen ces sist cetadla sce patties mene sts Sigalion ovigerum (15)
12. — Dorsal and ventral cirri foliaceous............... F. Phyllodocidae 13
13. —Five antennae, no occipital papilla; 4 pairs of tentaculan CirTiy eis vette lee LM EMenUR Esc ciaks Eumida sanguinea (13) —Four antennae and often an occipital papilla; 3-4 pairsiofitentacular cirri 55 aie aus gem uatayceie ce a ceee eee see ee mae eel) det es ee eh 14
14. —Setae present on third, or second and _ third, tentacullansepments() 6.01) fide ea fay ee tania at, cul i pear Sita ae rea ue Le 15 —Setae absent from all tentacular segments............... 0.0 cee eect eee 16
15. —Setae first present by second tentacular segment; rustyredimicolourya wens ne sees eee eee Genetyllis castanea ( 3) —Setae first present by third tentacular segment; colour otherwise, iiiyki van ederim gate Ne, Paranaitis (4 spp)
16. | —Pharynx with regular rows of papillae at sides of base, first 3 segments dusky, later segments with 3 dark
spots; dorsal cirri large and cordate anteriorly ..... Anattides longipes ( 3) —Pharynx with irregularly arranged papillae; body
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
17.
20.
21.
DR
23.
24.
25.
26.
27.
28.
31.
32.
33.
P. HUTCHINGS AND S.RAINER
pale to dark brown with dark intersegmental bands;
dorsal cirri almost semicircular................. Phyllodoce novaehollandiae —Prostomium with ventrolateral palps............... 20... 0c eee eee ee 18 — Prostomium:without palps! (0 0, sc8se Scie t gelesen sisse heels aes gleencre 51 —Palps biarticulate, with stout basal joint and smaller
Gistalhj orn teens tear neways Me waay cys tape ce een alien geen = Neste neciiay aay hous Sacileranas ran clave! ls Sa] sine ahaere 19 SAL PSISIMMP Lewy wapeierar velar enna eNO cence Nach p eco nece e eieeiin eye Sisal alocauecss Tetenere (eM e e e 35 —Compound setae absent; tentacular cirri two or fewer F. Pilargidae 20 —Compound setae present; tentacular cirri four pairs
OTETIVO TS ae ees Apt Nance sce seh smemah ear aflis Uanharie) am Petco a teal lee 51 Sul eoy vedios Uo RTO RBI Saltch Mciracte~ amen etae eng be 21 —Prostomium with three antennae; pharynx muscular Sigambra parva — Prostomium with two antennae; pharynx epithelial . Plargzs sp. —Jaws, if present, usually styliform, denticles absent;
tentacular cirri often jointed................... F. Hesionidae 22 —Two toothed jaws present and often horny denticles ;
tentacular cirrismooth....................... F. Nereidae 25 FO Mapalns tenctacUlan GME eigenen a cine ccaile wie esi os. cllaiese « shaceee Gl sere pete al eee eaeneleucee 23 SSIHIphtipalrsitentaculary Cai. ctr...) are oreut ty sree ksi aacla ue one UA hie © eiaeey See eae 24 —Two anterolateral antennae and one median
antenna, notosetae 1/2 capillaries............... Ophiodromus cf. agilzs —Two anterolateral antennae only; notosetae present
OlgabSem twa clete euens wis aae a nual et ousl Tats Gy, Nerimyra sp. —Notopodium reduced, with few capillary setae ..... Gyptzs sp. —Notopodium not reduced, with numerous capillaries. cf. Gyptis sp. —Anterior apodous segment absent behind
peristomium ; antennae absent; paragnaths absent.. Micronerezs sp. —Anterior apodous segment present behind
peristomium; two antennae present; paragnaths
PREsentvOr abseEMtbys «sis sien ee eames anid inamspraieuarsemt tee cae liane de oon at cbrsieuelepaitelay aero tenuis 26 —Chitinous paragnaths entirely absent; a transverse
series of glandular ridges at base of anterior
PATA POA welen wg.) sie, A els a ea ee ee IE NS, ata a ree Australonerets ehlersi —Chitinous paragnaths present; ventral glandular
MIO PEStADSEM CH Loe caine actin ee edeys eesorsh ye) Stee hac ncn eee com oben Sra Uma Bnah Ah teria ass 27 —Paragnaths all separate andconical ................ 0.0... c ee eee eee eee 28 — Paragnaths include pectinate or transverse bars..................0--- eee eeee 34
—Chitinous paragnaths present on both basal and ma Xd ary PUM GS 2, 2 2c sak aulsomaitayen wenpe ouianedamomanerte Sener dee Bien eoaae IevecleRtentaletay ssid sala A 29 —Chitinous paragnaths present on maxillary ring only .
—Falcigers present in posterior notopodia, few setae; paragnaths few, absent on I, V and VI, II-1, III +IV- ELV ANY) OereAY) 0 0 Et apenas te eiS, elie yee otter Sener ansae
—Falcigers absent in posterior notopodia...........
—Notopodial lobe very large on both sides of dorsal cirrus in median and posterior segments ..........
—Notopodial lobe not particularly enlarged
— Areas I and V of pharynx lacking paragnaths .......
— Areas I and V of pharynx with paragnaths
—Paragnaths of area I-2, and area VI-3 cones in in- ventedttrian glenn. eae atk era caer: oe — Paragnaths of area I-4 in triangle, and areas V + VIII — continuous band of cones 4-6 deep............
—Prostomium deeply cleft between antennae; dorsal cirri long; no simple neuropodial falcigers ........
Ceratonerets 33 Nerets postdoniae Neanthes 30
Neanthes oxypoda
Fae oie aoe eset Nena ORE 31
N. vaala
N. cricognatha
C. mirabilis
37
(13)
( 3)
( 3)
(13)
(13)
( 9)
(13)
(13)
(13)
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
38 ESTUARINE POLYCHAETES
—Prostomium not cleft; dorsal cirri short; one large
simple falciger in posterior neuropodia........... C. erythraeensts (18) 34. —Areas IV, VII + VIII of pharynx with cones and pectinate paragnaths, posterior notopodia with 2-3 homogomph falcigers........................ Platynerets dumerili antipoda (13) —Area VI of pharynx with continuous transverse series of cones extending across V and VI.............. Perinereis cf. brevicirris ( 9) 35. —Barrel-shaped proventriculus posterior to pharynx, pharyngeal armature variable.................. F. Syllidae 36 — Proventriculus absent, four or more pairs of jaws.............-...-0 ee eee eee 56 36. —Ventral cirri absent; dorsal cirri not annulated..... Autolytus sp. —Ventral cirri distinct, dorsal cirri annulated or SITLO OER PRG eS AHN caste il hoo Pala cae PNG ne BEAM ea Cte eB eae eeca bare COUR 37 37. | —Palps separate; dorsal cirrijointed ........... 0.0... eee ee eee 38 —Palps fused basally or for at least half their length; dorsalicirrismoothy cence epee uaa aie abana sia ue aarnsy eater ittama hey sae ee nev a eee eat 43 38. —Setae few, simple, enlarged, with 2 teeth, terminal One \bifid ei oa ee lay ce Meal eas ny eel ick hed ins aia Haplosyllis spongicola ( 3) —Setae mainly or entirely compound............... 0... ee eee eee 39 39. — Mainly compound setae anteriorly, then include a few large simple setae with Y-shaped prongs .......... Syllis graczles ( 3) = Onlyicompound'setaejpresenty os.) caiieie eise sl let et aici ieee a ern 40 40. — Blades of some superior setae much longer than the rest; dorsal cirri with 10-18 joints............... Langerhansza cornuta ( 3) —Blades of setae decrease evenly in length, dorsal cirri Otherwise! seis Ciena, Wate Voce rah ecole Steet iea yak Typosylles 41 41. —Setae unidentate or minutely bidentate, dorsal cirri stout with 8-12 joints ........................ T. armillaris ( 3) —Setae strongly bidentate; dorsal cirri otherwise..................- eee eee 42 42. —Dorsal cirri short, with 7-12 joints............... T. cf. hyalina ( 3) —Dorsal cirri with more than 20 joints............. T. variegata (13) 43. —Palps fused basally; dorsal cirri usually smooth. .................-200 eee ees 44 —Palps fused for at least half their length; dorsal cirri EJretolol a ohh eM eee ite elon amas dc TaRSR TEENS rata COs eA eM MENG Mts at Rn bien aiaate Gg 45 44. —Pharynx unarmed; ventral cirri longer than setigerous lobes; acicula of anterior parapodia enlarged, knobbed.......................... Streptosyllis sp.
—Pharynx with a semicircle of recurved teeth; ventral cirri not longer than setigerous lobes; acicula not
Knob bed eet Si a He ea a ga ariieea Odontosyllis sp. 45. —Dorsal cirri papilliform; 1 pair of rudimentary tentacular Ginnie is See a eS mittee hat) Exogone 46 —Dorsal cirri flask-shaped; 1 pair of tentacular cirri .. Sphaerosyllzs 48 46. —Dorsal cirrus present on setiger 2, single superior compound seta with dagger-like blade ........... Exogone sp. —Dorsal cirrus absent on setiger 2; superior compound setaeidifferent fromrest):. x y.ai Ppa Misses. eon taeevn ey mice ssa rsteis| ou oe ietl Starman 47 47. —Superior compound setae with swollen shaft-head and broad triangular blade....................... E. heterosetosa ( 3) —Three to five superior compound setae with dagger- like blades, c(i sic apis eile evehectia tera etch snesctisia art E. cf. gemmifera (13) 48. —Body surface and parapodia covered with minute Papillaey rs ire iiss Mirae ied eee eM Miinegs eis 2 fa Gra, ey i ed a at Cou NA canoe ae aa 49 — Body surface and parapodia without papillae ....................... 000s eee 50 49. —Dorsal cirrus on setiger2....................-.. S. cf. semzverrucosa (18) —Dorsal cirrus absent from setiger2 .............. Sphaerosyllis sp.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
50.
51.
Be
53.
54.
55.
Bite
59.
60.
61.
62.
63.
64.
P. HUTCHINGS AND S.RAINER
—Dorsal cirrus on setiger2....... —Dorsal cirrus absent from setiger 2
—Body papillose, head indistinct; pharynx unarmed . .
—Body smooth, head distinct; pharynx with two or {MOS OLUTION o bg oe odd oto sic obiclod olbipiaa 6b o dla Udo pololmeolo edo dia og o/6\c 52
— Peristomium with parapodia and setae — Peristomium without parapodia or setae
—Prostomium pentagonal, body parapodia with lamellae.......
square in section;
—Prostomium a pointed cone, body circular in section; parapodia-without lamellae’ ssn 2 sas ci lais wees sie cianejoieie ene aus ee Geena ene weer 56
—Prostomium produced anteriorly ;
—Prostomium not produced anteriorly; setae short
—Branchiae recurved, from setiger
long flowing setae.
4; pharynx with 22
longitudinal rows of papillae and single median
Papilla yar e eee wien oes —Branchiae recurved, from setiger
5; pharynx with 20
longitudinal rows of papillae and no single median
Papillary Wee ssa cs cviely ce sake soatets
—Pharynx with four horny jaws; and body not divided into regions
parapodia all alike
—Pharynx with a pair of toothed jaws and a circle of denticles; body divided into different regions ......
—Branchiae non-retractile, simple; pharyngeal papillae of two types, one with distal flange........ —Branchiae retractile, branched; pharyngeal papillae
of two types, without distal flange
— Body divided into three regions, parapodia biramous
after setiger 34..............
— Body divided into three regions, parapodia biramous
after setiger30..............
—Dorsal cirri and antennae present
—Dorsal cirri absent or rudimentary; antennae usually
ADSEN tae y votes atee a eos
—Maxillae of four or five paired plates, plates III and IV fused on right side; antennae variable; branchiae usuallyspresemt,: A. cls ciinct Mua pee ics a) afro aces decaf le el cegielsaemed na aan iv oh lal duasuee cu paancna sir 61
—Mazxillae of numerous small elements in two or four
longitudinal series, two antennae
palps; branchiae absent .......
and two cylindrical
— One to five antennae, without ringed ceratophores . .
—Seven antennae, posterior five ceratophores ...............
with long, ringed
— One antenna; branchiae and tentacular cirri absent
—Five antennae; branchiae present
—Tentacular cirri absent; anterior margin of prostomium bilobed; comb setae absent, compound setae spinigerous, acicular setae bidentate ........
—Tentacular cirri present.......
—Anterior margin of prostomium deeply notched, antennae deeply jointed ; branchiae begin on setigers 3-8; comb setae present, compound setae falcigerous,
acicular setae tridentate.......
Sphaerosyllis sp. S. sublaevis
F. Sphaerodoridae- Sphaerodoridium sp.
N. australiensis
N. inornata
F. Glyceridae 57
F. Goniadidae 58
Glycera tridactyla*
G. americana
Glycinde armigera
Glycinde sp.
F. Dorvilleidae 67 F. Eunicidae 62 F. Onuphidae 65 Nematonerets cf. unicornis Sey tesa Ato eae «AIO at mR eRe a 63 Marphysa sanguinea
Eunice 64 E. australis
*Glycera convoluta Keferstein is synonymous with G. tridactyla Schmarda
39
( 3)
(16)
(16)
(16)
( 3) (13)
( 9)
( 3)
(13)
(18)
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
40 ESTUARINE POLYCHAETES
—Anterior margin of prostomium shallowly notched, antennae nearly smooth; branchiae from setiger 3; comb setae present, compound setae falcigerous,
acicular setae tridentate...................... E. wttata ( 3) 65. —Branchial filaments arranged spirally, from setiger 5, pseudocompound hooks unidentate or with secondary toothivery weaken Vmnien gules ee vests si Seaways. letlea ie Diopatra cf. neapolitana ( 3) —Branchial filaments arranged spirally, from setiger 5; pseudocompound hooks bidentate .............. Diopatra sp. 66. —Prostomium conical, body elongate; hooded hooks jointed in first few parapodia, later hooks simple; body reddish or orange....................--. Lumbrineris latreill (13)
—Prostomium conical, body elongate; hooded hooks jointed in first few parapodia, later hooks simple;
body preemie pai Meena) ah esi entire ee ccmeaee teensy rn ceacnls Lumbrineris sp. 67. —Parapodia uniramous with ovoid dorsal cirri, no cirrophores or notoacicula.................... Protodoruvillea sp. — Parapodia sub-biramous with dorsal cirri mounted on ceratophores containing notoacicula............. Dorvillea australienszs (1)
Polychaeta Sedentaria 68. —Head not greatly modified; prostomium well developed; peristomium sometimes with pair of palps.....................+--- 69 —Head modified by development of frilly membrane, buccal tentacles or a branchial crown; prostomium
Oftensreduced yer Wye ce em ily a ete vs glos fate coves palaaacniaetse Sh Mr edieteeh Mays Stas 155 69. — Buccal segment with palps or tentacles ............ 0... eee eee ee 70 —Buccal segment without food gathering appendages ......................... 124 70. —Buccal segment with pair of adhesive palps (often broken off) or several grooved tentacles ............ 0.0... c cee eee 71 —Buccal segments with tentacles retractile into the TOUCH Sig lets er seer NG ee tector as F. Ampharetidae 160 71. —Hooded hooks present in posterior setigers; well developed parapodiiay yee) Mos ie Cane Teun ses eaves elo) ales hropaialegelienteaianauceneae anne 72 —Hooded hooks entirely absent; parapodia often poorlydeveloped eins Vii teemitec lets ay eueo tee nen eco ay nia thaws cilane sue Votieunite musi en Roane lit 108 72. —Head not flattened, branchiae often present; palps grooved ; posterior notosetae hooded hooks........ F. Spionidae 73 —Head flattened and spade-shaped; palps papillose, notosetae hooded hooks from setiger 9............ F. Magelonidae 106 73. —Setiger 5 with strongly modified setae .............. 0.02. cee eee ee eee 74 —Setiger 5 without modified setae ............ 0.0... eee ee 85 74. —Branchiae first present posterior tosetiger5 ............ 0... e eee eee eee eee 75 —Branchiae first present anterior to setiger5........... 00-0. eee eee eee 83 75. —Setiger 5 slightly to moderately modified parapodia.. | Pseudopolydora 76 —Setiger 5 greatly modified, with reduced parapodia.............-...--++-++--- 77 76. —Prostomium entire; neuropodial hooded hooks from Sebi pen Ora sary Hi ania ess) aye chen rae amen veren te Olen CO si P. paucibranchiata ( 2) —Prostomium incised; bidentate neuropodial hooded hooks fromysetiger 8 7.cy.0sey mee ee eee tr uieas P. kempi ( 2) 77. —Setiger 5 with spines of one type with or without companion capillarysetae .................... Polydora 78 —Setiger 5 with spines of two types................ Carazziella 82 78. —Setiger 5 with brush-tipped companion setae; specialized posterior notosetae absent............ P. penicillata (13) —Setiger 5 without brush-tipped companion setae; specialized posterior notosetae present or absent ..........-----22+-e seer ee eee 79
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
80.
81.
82.
84.
85.
86.
87.
88.
90.
91.
P. HUTCHINGS AND S. RAINER 4] — Hooded hooks with constriction on shaft.......... P. haswelli ( 2) — Hooded hooks without constriction on shaft ............ 0... e ee eee eee ee 80 —Posterior notopodial spines form dense packets of needles; gizzard absent; major spines of setiger 5 with weak subterminal swelling; eyes absent........... P. flava ( 2) — Posterior notopodial spines absent ................ 0... e eee ee eee 81 —Gizzard present internally in setigers 18-19, seen externally as a swelling; major spines of setiger 5 with subterminal boss; eyes present ................. P. socialis ( 2) —Gizzard absent; major spines of setiger 5 falcate, simple seyesjabsentannyeiys acne ara nsie aera Polydora sp. —Hooded hooks from setiger 7; superior dorsal fascicle of notosetae with fimbriated setae............... C. hirsutiseta ( 2) —Hooded hooks from setiger 8; superior dorsal fascicle of notosetae with simple setae.................. C. victoriensis ( 2) —Major spines of setiger 5 of two types, first with expanded ends bearing bristles, second simple, fal Cate pret crane ne seen td seine. Litas UMM nad fu Boccardza chilensis ( 2) — Major spines of setiger 5 of one type, simple, falcate, with smaller companion setae.................. Boccardtella 84 —Branchiae from setiger 2 onwards except for setigers 4 ATID erences cyte sarae yume mo ma ANE eee ee ee bore B. bihamata ( 2) —Branchiae from setiger 2 onwards, on setiger 5 may be RU GIMEM GARY) Cy spaleesitee ests srodunanaia ea al sea satay Mita B. limnicola ( 2) —Prostomium distally pointed............ 0.0... cee eee ee eee 86 —Prostomium not distally pointed, with lateral or frontal horns, anterior margin broadly rounded or aS Ui Bias ve echt vie a RE ATEGe Rae OU EntEE Sn Ook PEL RMD eh RR erate ES TEE UE rey 90 —Branchiae beginning on setiger 1, continuing almost LOMMOStETION.EMGs silo o hla faracs eatiesele) oo) seinda lara sis os Australospio trifida ( 2) —Branchiae beginning on setiger 2, continuing over Vatiablemumiben of setigers)< wists ncn scree craciee acl sche eon or a ev etien clare lel arate eaten 87 —Branchiae completely free from dorsal lamellae, albsentjposterionlys s)sarinniee aie ene ocak Aonzdes oxycephala ( 2) —Branchiae fused to dorsal lamellae and continuing to eEndiofibodye sii sus ley Sees toca ais silane tee sustal Scolelepis 88 —Notosetae present on setiger 1, neuropodial hooks quadnidentate Yee isin) swaenae aula Gets ld heats S. precirriseta ( 2) —Notosetae absent on setiger 1, neuropodial hooks Otherwise; iscsi) i cake ole hence tae Rew aa od teen tear ne eau lelic sn Dut SAM es NM air SINS 89 —Hooded hooks multidentate, from setiger 14-15, in Meuropodia only. oi. es ee oe aol ein elel es S. towra ( 2) —Hooded hooks bidentate, from setiger 24, probably onlyin neuropodia..:...................04.- S. vexzllatus (as Pseudomalacoceros vexillatus) (13) —Prostomium with lateral or frontal horns...............-0-- 0s esse cette tees 91 —Prostomium without lateral or frontal horns .................... 0000s eee eee 94 —Branchiae from setiger]...................... Malacoceros 92 —Branchiae from setiger 2...................... Rhynchosfio —Bidentate ventral hooded hooks from setiger 22... .. M. divisus (13) —Tridentate ventral hooded hooks from setiger 11 or ACT Pee Ley ays ese ee ect eee ie Ma MeN cTectesie MRCP ENE es ntd uate abiuers -aitaplur ae acta ac arehial a) # Sonata ia 93 —Body with dark brown pigment; tridentate ventral hooded hooks from setiger 25 .................. M. tripartitus ( 2)
—Body colourless; tridentate ventral hooded hooks from setiger 11........
Malacoceros sp.
Proc. Linn. Soc. N.S.W., 104
(1), (1979) 1980
42
Wee
why
96.
97.
98.
99.
100.
101.
102.
103.
104.
105.
106.
107.
108.
109.
110.
111.
112.
—Neuropodial
ESTUARINE POLYCHAETES
—Branchiae limited to middle and posterior setigers
except for one pair on setiger 2inmales ..........
—Branchiae from setiger 1 or 2
—Branchiae concentrated in setigers 1-22 —Branchiae present over most of body
—Branchiae from setiger 1 —Branchiae from setiger2......................
—Branchiae otherwise
—Branchiae all pinnate —Branchiae both pinnate and cirriform
—Three pairs of pinnate branchiae; dorsal ridge across SECIBerul reese —Kighteen to 22 pairs of cirriform branchiae; no dorsal ridge on setiger 1 ..
=Branchiae.all cinnformen eee eee
Prionospio
Paraprionospio sp.
Orthoprionospio cirriformia
P. ctrrifera
Pie aoe eee er ry AUB Re Cat cet ea Pe Reo aI PL MOE a ey i 99
— Three pairs of pinnate branchiae; low dorsal crest on Seliger Mase ee —Four pairs of pinnate branchiae, dorsal crests absent .
—First and third pairs of branchiae pinnate (pinnules
sparse) , second and fourth pairs cirriform.........
—Branchiae otherwise
—First three pairs of branchiae cirriform, fourth pair pinnatey ae ade —First and fourth pairs of branchiae pinnate, second and third pairs cirriform
— Dorsal transverse crest on setiger 7 only........... — Dorsal transverse crests from setiger 7 to about 30...
—Branchiae from setiger]...................... —Branchiae from setiger2..................-.-..
—Neuropodial hooded hooks from setiger 10-11,
bidentate; body colourless ....................
hooded hooks
from setiger 9-11,
tridentate; anterior body brown pigmented .......
—Hooded hooks from setiger 9, tridentate (MF:2),
accompanied by single short hooded hook per fascicle
—Hooded hook from
multidentate
setiger 9, bidentate or
—Hooded hooks multidentate, all hooks within a
fascicleisimilarinisizey sa ee ie
—Hooded hooks bidentate (MF:1), all hooks within a
fascicleisimilarminisize..) ae A eee
—Long filamentous branchiae at least on anterior
segments; parapodia reduced toridges...........
— Two large grooved tentacular filaments —Several grooved tentacular filaments
— Capillary setae only present —Acicular hooks and capillaries present
just above notosetae
dorsalline .......
—Branchiae not long and filamentous; parapodia not imithe form Of Tid ges i. yee del uname apace anaes ec cena moe teen at sae ote Nee
—Branchial filaments in middle of body arise laterally
—Branchial filaments in middle of body arise from mid
—Acicular setae of posterior noto- and neuropodia
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
P. aucklandica P. multipinnulata
P. paucipinnulata
P. fallax P. multicristata
Spio Laonice sp.
S. filicornas
S. pacifica
Magelona dakini
Magelona sp.
Magelona sp.
F. Cirratulidae
T. marioni
T. dorsobranchiata
100°
101
102
103
105
107
109
( 3)
( 2) ( 2)
( 2)
( 6) (13)
( 3) ( 2)
(13)
( 3) ( 3)
113.
114.
115.
116.
117.
118.
119.
120.
121.
122.
123.
124.
125.
P. HUTCHINGS AND S. RAINER
continuous to form two lateral arcs, on either side of body; acicular hooks present in only last 40 setigers, increasing rapidly to 5-6 per fascicle plus 5-6
Capillaniess apie cis cree enn Chaetozone sp. —Posterior noto- and neurosetae remain in separate
uma Tes pee te snes tcuisnararat ch ecchlansbesal ass aberemelonss Caullerzella 113 —Neuropodial hooks from setiger3............... Caulleriella bioculatus —Neuropodial hooks from setiger 3 or later... 1.1.2.0... 0.0 cece eee eee eee ees 114 —Neuropodial hooks from setiger 4, bidentate....... Caulleriella tricapillata —Neuropodial hooks from setiger 31 or later, bidentate........................ 115 —Neuropodial hooks from setiger 31, minutely
multidentate, accompanied by capillaries; single
acicular seta present in posterior segments; elliptical
EV EIS POE ae eet sscysy nye oseiioy seisy ahs naerter aieereyssas fevied cietce sas Caullerzella sp. —Neuropodial hooks from setiger 34, multidentate by
setiger 71, accompanied by capillaries; simple
acicular hooks posteriorly, sigmoid anteriorly; pair of
darkseyeispotss eye iiacysn a) als ie sunce eee o syetnoss ses its Caullerzella sp. —Branchial filaments arise om same segment as
tentacullanicirnie ssnyacies 4) cisele es ecm aiameeale als Cirratulus 117 —Branchial filaments arise on all segments anterior to
tentacularteimniy aii ut ep pepe deme chad a Cirriformia 119 —Tentacular filaments arise above setigers4-6....... Cirratulus chrysoderma —Tentacular filaments arise otherwise. ......... 20... 0. eee eee eee 118 —Tentacular filaments arise above setigers 6-10;
acicular setae absent, capillaries throughout;
anterior heavily pigmented.................... Cirratulus nuchalis —Tentacular filaments arise above setiger 3; sigmoid
hooks and capillaries present .................. Cirratulus sp. —Tentacular filaments arise above setigers 3-4....... Cirriformia capensis —Tentacular filaments arise above setigers 4-6 0r5-7............... 0000000005: 120 —Tentacular filaments arise above setigers 4-6;
capillaries throughout, sigmoid hooks by setiger12.. Crrréformza filigera —Tentacular filaments above setiger 4-6 or 5-7;
capillaries throughout, sigmoid hooks from setiger 50 Cirriformia tentaculata —Noto- and neuropodia of anterior parapodia well
developed, setiger 1 with elongate setae directed
POT WAT ASI tsetse ccs ueik sea cote e Cena psiesn or gual cioctad F. Trochochaetidae-
Poecilochaetus serpens
—Anterior setigers uniramous, neuropodia lacking;
posterior setigers biramous with neurosetae as minute
(UHDYSNT OV Sey a ee URN cNIneA amnion GiciAtan Dl eee Sonia nor erase Ole F. Chaetopteridae 122 — Middle region with bilobed notopodia; no tentacular
Pel 9 Welt AU PS ac EARN citcett Spee NB LS oO Spiochaetopterus sp. —Middle region with notopodia never bilobed; no
temtac ular Cir s) eee 255) .caleh seem eae amy ae istientcltaun sn ate besa calibte geal louis fantap lanes aie aiy tat Alle tate 123 —Body large; palps short; some notopodia of middle
region fused to form paddles; parchment tube ..... Chaetopterus variopedatus — Body small; palps large; notopodia of middle regions
never fused; fragile sandy tube; often living in dense
COLONIES sie Supe Reescis elas ae aici peet ee non eueee ale Mesochaetopterus minutus — entatecrestedshooks absent sisi e ee enciey lei neti ee) ol We clievients) aii ilcliclte rele eetistie eile 125 —Dentate crested hooks present'in posterior segments if
INCOXe GEMAb VEC! eb 656 .6.1B'0.4'6 oibLg Ao\ p10. 0/0'0'4. 0°o ol OU'E OlS Beeb Ole GC Oploa'd 06.000. Olan '5 O.0!0 138 —Capillary setae crenulate ..................... F. Orbiniidae 126 = C@apillanysetae smoothiarniy. cite ie rye torres eciei bene acl ner es ete ele) eee) test 132
Proc. LINN.
Soc. N.S.W., 104 (1), (1979)
43
( 3)
(13)
( 3)
( 5)
( 3)
( 3) ( 3)
( 3)
( 3)
( 3)
1980
44 ESTUARINE POLYCHAETES
126. —Prostomium round; change from thorax to abdomen about setigen 280.) 72 Sige ces cute ire sateen aes a
—Prostomium pointed ; thorax of less than 23 setigers ....................-5---.
127. | —Thorax with several foot papillae and many stomach Papillae yy ie eee ise ee are el aues tile lal ah ok oer eras eas — Thorax with 1-2 foot papillae, no stomach papillae.......................... 128. —No anterior row of hooks among thoracic neurosetae . —Anterior row or rows of hooks among thoracic NEUFOSELAE 7h eva sti cct occult Halts yo elionead AaNteN Sea boy arate ueiiey « Rearenaan Ween oe gee uNse7 es omen 129. — Thorax of 19-22 setigers; branchiae from setigers 8- A Ca ee I eile cara NG ie se oes acrae OUR Sa eae Reh — Thorax of 14-15 setigers; branchiae from setigers 15- UG ye sae h ieee peat eatertapninlae sekim ee Isyattdelieigen cds suum 130. —Single row of slender hooks among thoracic MEUTOSEE AE). eo) Ue zagch su nae al aise een cee h cra uae rolcapa ate —Three to four rows of hooks among thoracic MEUNOSEL AEA cil n Ne nee Sees atu ses: ale Maan otnat a 131. — Thorax of 13-17 setigers; branchiae from setiger 9-11 — Thorax of 16-18 setigers; branchiae from setiger 12- Dai ti Masala co tale cy Sane oan ana tMiou yin ie erere Te Ese Ie oc tm Be 132. —A single long filiform branchia arising from dorsum OSEtIPETs2 OT. See iat ieee ced a neh eae a ote as
—Branchiae, if present, not single and median...................0--e see eeeee
133. —Capillaries winged in anterior segments; median antenna may be present ...................... —Capillaries not winged; median antenna never
[eices{lelee al oMteee eeuc of beni Me mao ncn bleep olMiog nino tid amie sa thats Midie ian a bina obo g'efh bo
134. —Prostomium with median dorsal antenna; specialized setae, 4 curved unidentate hooks among capillaries of posterior neuropodia ......................0-.
—Prostomium without an antenna; specialized setae among capillaries of posterior neuropodia.........
135. —Prostomium a tapered cone; body fusiform, often groovediventrally eer oe bos | ates oetien: —Prostomium notched or lobed; body swollen anteriorly but not grooved ventrally .............
136. —Ventral groove present; branchiae present from setiger 2 for about 24-25 setigers; eye spots between parapodium of middle segments................
—Ventral groove present, branchiae absent; eye spots El of oin ast its es Vaile a abaiolee be fokata oy ale tupes oracle
137. — Body arenicoliform; posterior parapodia lack ventral and dorsal cirri; branchiae absent ..:...........
—Body arenicoliform; posterior parapodia with both
ventral and dorsal cirri; branchiae absent.........
138. — Body resembles an earthworm; dentate crested hooks withthoods) ise al os eaten anh 1a ayes Ol Feit ran areal —Body does not resemble an earthworm; dentate
crested hooks without hoods .....................
139. —Genital setae present in both sexes on setigers 8-9; thorax/ofil Ojsetigersiy egy eis eee eel — Genital setae absent or only present in males; thorax
Of setigens 2s sry utenti Nave anaes al ay aia WLSsIonG aie arty LEM eer egy ennui eg erie
140. —Genital setae present in males; thorax of nine
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
Nainerts grubet australis
L. bifurcatus
L. normalis
Scoloplos (Scoloplos)
Scoloplos (Leodamas) johnstonez
Scoloplos (S.) simplex
Scoloplos (S.) sp.
F. Cossuridae-Cossura sp.
Aricidea fauveli
Paraonis sp.
F. Opheliidae
F. Scalibregmidae
Armandia intermedia
Polyophthalmus pictus
Hyboscolex dicranochaetus
Pseudoscalibregma sp.
F. Capitellidae
(4, 13) 127 (4, 13) 128 129 130 (4, 13) (4, 13) 131 ( 4) (4, 13) 133 134 135 ( 3) 136 137 ( 3) ( 3) (14) 139 149 (13) 140
141.
142.
143. 144. 145.
146.
147.
148.
149.
150.
151.
152.
153.
154.
155.
156.
P. HUTCHINGS AND S. RAINER
setigers; hooks thereafter............
— Body colourless; when preserved
capillaries in anterior thoracic setigers,
— Body with bright red pigment spots..............
—Ten thoracic setigers, with capillaries in setigers 1-4
and hooks in setigers 5-10.....
—Morejthaniten' thoracic setigersjisn asses saci a ae ce dea eee ae
el eVveEnbENOTACleisebiGensnn hi Uian clan mics Maayncel ential eniie sia thw sync usatiscura tas tye leve = Morethanyl lithoracicisetigersiny «a says ise cic tees oe ieee nes eeiese eo le oe cee
— Thoracic setigers all capillaries .
— Thoracic setigers with capillaries and hooded hooks
—Neurosetae present
—Neurosetae absent on first thoracic setiger
—Eyes embedded in segment 1; globular branchiae
on first thoracic setiger ........
present on posterior abdominal setigers...........
—Elliptical eye spots on segment 1; often purple in
colour; branchiae absent....................-.
— Thoracic setigers 1- — Thoracic setigers 1-
—Twelve thoracic setigers; abdomen ends in anal
5 with capillaries, 6-11 with hooks 6 with capillaries, 7-11 with hooks
plate; branchiae absent......................
— Thirteen thoracic setigers; no anal plate; retractile
branchiae
—Middle setigers greatly elongated, never annulated ;
branchiae rare...
—Middle setigers not greatly elongated but annulated ;
branchiae present.
—Cephalic plate absent; cephalic ridge well marked; pygidium petaloid with centralanus.............
—Cephalic plate well defined, surrounded by raised margin; pygidium variable
— Pygidium as a slanting plate with a dorsal anus above
it, anal cirri absent
—Pygidium encircled by anal cirri with anus sunken in EV IADR OVS ei ee hi React easy ariaherc by oly ace melee eo sels Ale bars BRCM n a esa Water e Wala gum veatae winec. cen acc
—Neurosetae of setigers 1-3, four dentate crested hooks similar to subsequent ones —Neurosetae of setigers 1-3, four acicular setae,
different from subsequent ones.................
—Setigers 1-3 with simple smooth acicular neurosetae; 2 achaetous pre-anal segments; anal funnel with 27-
29 equal cirri plus one longer ..................
—Setigers 1-3 with simple smooth acicular neurosetae ; five achaetous pre-anal segments; anal funnel with
allicinnilequalhimisizemiar nice iii ror aac nl
—Branchiae from setiger 20, branchiferous segments
with two annuli; animal small not exceeding 40mm .
—Branchiae from setiger 7-17, branchiferous segments
with five annuli; animal large..................
—Head with frilled food gathering membrane; lives in
firm sandy tube . .
—Head without frilled food gathering membrane
ead withistoutsetae ian for neo
— Head without stout setae
Proc. Linn. Soc. N.S.W., 104 (1)
Capitella 141 Medea seas Penmaes essctemer amet 142 C. capitata Capitella sp. Mediomastus californiensis 143 144 148 Notomastus 145 SOI ELON ONION Manor eere 147 Notomastus sp. Baa bags Ira i ape ane ON BAe mae ae ects 146
N. torquatus
Notomastus sp.
Heteromastus filiformzs Barantolla lepte
Scyphoproctus djiboutienses
Dasybranchus sp. F. Maldanidae 150 F. Arenicolidae 154 Petaloproctus sp. SISO Ico eR Ee TERE roa rAaT 151 Asychis sp. 152 Axiothella sp. Euclymene 153 E. trinalis Euclymene sp. Branchiomaldane sp. Arenicola bombayensis F. Oweniidae-Owenza fusiformis ai Cee rowel coat eNt Wot eb reamed 156 F. Flabelligeridae 157 Hit as Hen AS ne aoh Sta ha 159
45
( 3)
(13)
(13) (13) (13)
( 3)
(11)
(13)
( 3)
, (1979) 1980
46 ESTUARINE POLYCHAETES
157. —Neurosetae simple hooks; branchiae filamentous, arise in single marginal row from cephalichood .... = Pherusasp. —Neurosetae not simple hooks; branchiae filamentous or cirriform(\arisejin'2(or, MOre|TOWS))s als. elie ee 158
158. —Neurosetae annulated with bent or minutely hooked tips; a few stout branchiae in 2 distinct groups ..... Diplocirrus sp.
—Neurosetae stout compound hooks; numerous fine branchiae in several irregular rows .............. Flabelligera sp.
159. —Head with soft tentacles for deposit feeding; branchiaempresemt ss.) 25h Syste ees ae esi orsitet elec aie fe cen ania baleen -o aratrn er atte are ptr ee 160
—Head with branchial crown for suspension feeding ; segmental }branchiae/absemty. 6 she ei ars aiccssrnicciacein upset ten ree ee eee ents eee 179
160. —Tentacles retractile into mouth, either grooved or Papillose ye) Rye het eMon i amote coh enseetelatiags F. Ampharetidae 161
—Tentacles not retractile into mouth, grooved, never Papillose secs richmn AS Wauwut crc Aaara ECA my caes lala what earls eget aver Aer wt ema dis anon bere 164
161. —Stout postbranchial hooks present; paleae absent; buccal tentaclessmooth ...................... Isolda pulchella ( 3)
162. —Buccal tentacles papillose..................... Pseudoamphicters papillosa (12) —Bucealitentacles smooeh 5.72 /51.05 yrs ye Sia ites ict i cite ete ee talie alee mene aleeete 163
163. —Glandular ridges present; four pairs of smooth gills; 14 uncinigerous thoracic segments; body with green PISMENt Spots see sri ite seen eee ete epee aaa Amphicteis dalmatica (13) —Glandular ridges absent; three pairs of smooth gills; 12 uncinigerous thoracic segments .............. Samythella sp.
164. —Branchiae absent; tentacular lobe expanded ...... F. Terebellidae SF. Polycirrinae 165 —Branchiae present; tentacular lobe expanded or COMP ACC Meee Hii oneal aye yh UN cre ane ea Ceeebs aay usu ai Oe Ate ci ee OR 167
165. — Abdominal setae completely absent ............. Lyszlla 166 — Abdominal setae restricted to acicular notosetae.... Amaeana trilobata (12)
166. —Ten thoracic setigers; notosetae simple smooth capillaries, neurosetae absent.................. L. apheles (11)
—Nine to 12 thoracic setigers; notosetae barbed, Meurosetaejabsentar yay ieee cane ae L. pacifica (12)
167. —Tentacular lobe and peristomium compact........ F. Terebellidae 168 —KEither tentacular lobe or peristomium expanded.... _ F.. Trichobranchidae 178
168. —Branchiae simple filaments on setigers1-3.......... 0.0.00... e cee eee eee ee 169 —Branchiae branched or club shaped on setigers 1-3 ................000- 00 ees 174
169. —Uncini of first four rows, heavily chitinized long- handled hooks; following ones avicular........... Hadrachaeta aspeta (12) —Uncini all avicular with forwardly projecting bases ....................000--- 170
170. —Notosetae from segment 2 (first branchiferous) .... Streblosoma 171 —Notosetae from segment 3 (second branchiferous) ....................00-00- 172
171. —Twenty-five—29 pairs of notosetae; three pairs of branchiae forming continuous band across dorsum . . S. acymatum (13) — Twenty-one pairs of notosetae ; two pairs of branchiae with distinct median gap separating left and right hand! pairsye yey incre em hin tees cuekomouet Streblosoma sp.
172. —Notosetae from segment 3 (second branchiferous) ; UN cinitKomNsetiper saree eee cen Thelepus 173 —Notosetae from segment 3 (second branchiferous) ; UNGInipfromiIsetigerO4 Ae Eeiocn i riarece: Rhinothelepus lobatus (11)
173. —Notosetae terminate halfway along abdomen, uncini
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
P. HUTCHINGS AND S. RAINER
with terminal button; branchiae of numerous
fala rrvem siete here save See eee core eau cake Aaeaeatients T. setosus —Notosetae more than 20 pairs; uncini otherwise;
three pairs of branchiae with few branchial filaments
arising on horizontal ridges, filaments of first extend
Vatenalllyies eae its es sauce Vie Ase eb teases bas teehee Thelepus sp. 174. —Branchiae club-shaped; anterior uncini avicular but base produced backwards as long shaft........... Pista typha —Branchiae branched; all uncini similar, either avicular or pectiniform...... IBA RIGHT aM na ce ee tea oi Mee cd Soe 175 175. = Notosetae with; smoothitips <av..scs pees 6 lotro) Pesta) ey oho dba sien vemiautee idl elie ye 2) 8a ala sane 176 = INotosetaeswithidenticulatel tips) try oily cies ietieie) sesh ibis hacia cusa as = 177 176. — Lateral lobes absent on segments 2-4; uncini avicular Nicolea sp. —Lateral lobes present on segments 2-4; uncini PE CEITMITORIM Sy iio Sloe anny Matas rec ae aie ane at alias Loimia medusa 177. —Lateral lobes absent; notosetae from segment 3 (second branchiferous) ...................... Neoleprea sp. —Lateral lobes absent; notosetae from segment 4 (third branchiferous)................--.+.--- Terebella sp. 178. —Expanded tentacular lobe; compact peristomium ; single branchiae with stout trunk bearing four lamellatetlobesy Gaye is are, Sane as ce se chaeyerelied Se, autancalte Terebellides stroemz
—Compact tentacular lobe; peristomium forming conspicuous proboscis; branchiae simple filaments on SEEGERS Sigs ac ts) sels serene eh ey intial tase suse Mig Aa Artacamella dibranchiata
179. —Tube sandy or muddy; an operculum never present AMONG Tad Olesn site's, fae, cisco ca nyls wean sue=tyecmueicerey ey see F. Sabellidae 180 —Tube calcareous; stalked operculum usually present. __F. Serpulidae 183
180. —Thoracic neurosetae single row of long-shafted hooks; last few abdominal segments form a spoon- Shiapedicavityie nhs waka oc eau Mecn he kp eayctedeae rman Euchone sp. —Thoracic neurosetae avicular; posterior abdominal Sepmentsimotmodifiedy sy seo cesses vaecle ciap aly ae jaSeiss evastege nantes leet see eben m lalvara eulclnu arate 181
181. —Thoracic neurosetae avicular uncini with pick-axe setae; collar replaced by triangular projection ..... Amphiglena pacifica — Thoracic neurosetae avicular uncini without pick-axe SERIO. 3 o'b Wale Menor nly Gitl'o (op "nn cols lotalas ayaa oles: 6 olalov od trove pho ay aore in-arevorg 6 atauavarnaase 182
182. — Thoracic notosetae include subspathulatesetae...:. cf. Laonome sp. —Thoracic notosetae winged capillaries only; branchial radioles with stylodes; body with segmental
eye spots and irregular dark pigment............. Branchiomma ngromaculata 183. — Thorax of 3-4 segments; tube small, spirally coiled.. Janua 184 —Thorax of 5 or more segments; tube straight or SUMUMOU SW ee eee ate aa ssn eee een ee Mean MOU atlas a Sis ain teaivw ina) de ene ogee NW eRe MeN NC ele aa 185 184. — Tube with one to four longitudinal ridges; operculum with peripheral bilobed talon.................. J. (Dextospira) brasilienses —Tube with three longitudinal ridges; operculum GOMGAVE ty yee tii 9a Nanas Mae pace ean anarrarhts fille J. (Dexiospira) foraminosa 185. —Collar setae bayonet; operculum stalk slender,
wingless, operculum with basal funnel or fused radii
plus central crown of 10-20 equal horny spines with
lateralis pimalesiy cite ted cam ee alia ta Hydrozdes elegans * —Collar setae very short and fine; operculum winged,
operculum with three-four basal plates from base of
which arise nine moveable spines, inner smooth, outer
SIME Aeterna A Nel Sra 2s MUMBA Rr Galeolarza caes pitosa
*Hydroides norvegica Gunnerus is synonymous with H. elegans (Haswell)
47
( 3)
(12)
(12)
( 3)
(12)
(13)
(13)
(13)
(13)
( 3)
(17)
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
48
16.
17.
ESTUARINE POLYCHAETES
References
BENHAM, W. B., 1915. — Report on the Polychaeta obtained by the F.1I.S. ‘Endeavour’ on the coasts of New South Wales, Victoria, Tasmania and South Australia. Part 1. In H.C.Dannevig (ed.) Bzol. Res. Exped. F.I.S. Endeavour’ (Sydney: Ministry for Trade and Customs) 3 (4) : 171-237.
BLAKE, J. A., and KuDENov, J. D., 1978. — The Spionidae (Polychaeta) from Southeastern Australia with a revision of the genera of the family. Mem. Nat. Mus. Vic. 39: 171-281.
Day, J. H., 1967. — A monograph of the Polychaeta of South Africa. Vol. 1 and 2. London: British Museum (Nat. Hist.) .
——, 1977. — A review of the Australian and New Zealand Orbiniidae (Annelida — Polychaeta) zn Reish and Fauchald (eds), Essays on Polychaetous Annelids in Memory of Dr Olga Hartman (pp. 217-246). Los Angeles: Allan Hancock Foundation.
—, and Hutcuincs, P.A., 1979. — An annotated check-list of Australian and New Zealand Polychaeta, Archiannelida and Myzostomida. Rec. Aust. Mus. 32: 80-161.
EHLERS, E., 1907. - Neuseeliandische Annelidenl1. Abh. K. Ges. wiss. Gottingen (Math. — Phys. Kl.) n.F. 5(4): 1-31.
FAUCHALD, K., 1977. — The Polychaete Worms. Definitions and Keys to the Orders, Families and Genera. Natural History Museum of Los Angeles County, Science Series 28: 1-190.
Foster, N. M., 1971. — Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the fauna of Curagao and other Caribbean Islands. 36 (129) : 1-183.
HARTMAN, O., 1950. — Polychaetous annelids. Goniadidae, Glyceridae, Nephtyidae. Allan Hancock Pacif. Exped. 15: 1-181.
—— , 1954. — Australian Nereidae. Trans. R. Soc. S. Aust., 77: 1-41.
Hutcuincs, P.A., 1974. — Polychaeta of Wallis Lake, New South Wales. Proc. Linn. Soc. N.S. W.
98: 175-195.
——, 1977. — Terebelliform Polychaeta of the Families Ampharetidae, Terebellidae and Trichobranchidae from Australia, chiefly from Moreton Bay, Queensland. Rec. Aust. Mus. 31: 1-38.
——, and RalIner, S. F., (1979). — The Polychaete Fauna of Careel Bay, Pittwater, New South Wales, Australia. J. Nat. Hist. (Lond.) 13: 745-796.
KupeEnov, J. D., and Bake, J. A., 1978. — A review of the genera and species of the Scalibregmidae (Polychaeta) with descriptions of one new genus and three new species from Australia. J. Nat. Hist. (Lond.). 12: 427-444.
Monro, C. C. A., 1924. — On the Polychaeta collected by H.M.S. ‘Alert’ 1881-1882. Families Polynoidae, Sigalionidae and Eunicidae. J. Linn. Soc. Lond. (Zool.) 36: 37-64.
RAINER, S. F., and Hutcuincs, P. A., 1977. — Nephtyidae (Polychaeta; Errantia) from Australia. Rec. Aust. Mus. 31: 307-347.
STRAUGHAN, D., 1967. — Marine Serpulidae (Annelida: Polychaeta) of Eastern Queensland and New South Wales. Aust. J. Zool., 15: 201-261.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
The Vegetation on two Podzols on the Hornsby Plateau, Sydney
ROBIN A. BUCHANAN and G. S. HUMPHREYS
BUCHANAN, R. A., & HumpuHReEys, G. S. The vegetation on two podzols on the Hornsby Plateau, Sydney. Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980:49-71
Podzol soils on the upland sandstone plateaux in the Sydney Basin have received little attention and a characteristic vegetation on these soils has not been reported previously.
Two sites on the Hornsby Plateau are selected for detailed description of the understorey vegetation on these soils and the type of vegetative changes that occur across a podzol/non-podzol soil boundary. The two podzols are in topographically different situations, one on the plateau surface and one in a gully, but like all podzols in the area they occur in sand deposits which are usually present in areas of low slope. The vegetation on the podzols in both cases is similar and typical of that on most podzols of the dissected sandstone plateaux around Sydney.
Ceratopetalum gummiferum, because of its growth form and foliage, is usually the most conspicuous species on the podzols, even where it is not abundant. Other common species are Banksza serrata, Xanthorrhoea arborea, Xylomelum pyriforme and to a lesser extent Gompholobium latifolium and Ricinocarpos pinifolius. Plants growing on podzols usually have a wide habitat range or are species of the upper and middle gully slopes. Species typical of the plateau surface are usually absent or infrequent, even where the podzol immediately adjoins such a community. A combination of physiography (low slope depositional areas) and floristics can be used to locate these soils.
The woodland or open-forest on podzols is usually dominated by the widespread
species Angophora costata and Eucalyptus gummifera and is usually taller than on surrounding soils. Eucalyptus haemastoma is extremely rare on podzols. : Robin A. Buchanan, 22 Alicia Road, Mt Kuring-gaz, Australia 2080 (formerly School of Biological Sciences, Macquarie University) and G. S. Humphreys, School of Earth Sciences, Macquarie University, North Ryde, Australia 2113; manuscript received 2 July 1979, accepted in revised form 19 September 1979.
INTRODUCTION
A complex mosaic of vegetation is present on the deeply dissected, species-rich sandstone plateaux around Sydney. Because of this deep dissection, the flora can be grouped according to broad topographic situation with species most common on the plateau tops, on the slopes and in the gullies. The distribution of structural forms of the vegetation also follows a general topographic pattern, with variations in both species composition and structure influenced by aspect and climate (Pidgeon, 1937, 1938, 1941; Burrough, Brown and Morris, 1977).
However, many marked variations in the vegetation occur over distances of only a few metres and cannot be interpreted in terms of aspect, climate or macro- topography. Differences, although often minor, in_ soil properties and microtopography provide the best explanations of these changes. The rapid change in soil characteristics between a podzol and the neighbouring soil is a dramatic example of the sudden floristic change which can accompany a change in soil type.
Podzols in the Sydney region of New South Wales are well-known from the coastal dune sands and Nepean terraces (Burges and Drover, 1953; Walker and Hawkins, 1957; Walker, 1960). The podzols associated with the sandstone plateaux are at a higher topographic level and form a distinct third group, which previously has been given no more than cursory attention (Walker, 1960; Hamilton, 1976; Forster et al., IaD)e
An initial field survey on the sandstone plateau of Lambert Peninsula in Ku-ring- gai Chase National Park (Buchanan, 1980) showed that podzol soils occur in areas of
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
50 VEGETATION ON PODZOLS
quartz sand deposits. These deposits are 1-4 m thick and less than 5 ha in area and have accumulated in places of low slope either near the plateau surface or in gullies. The dark organically-stained upper-surface layer of the sand, 10-20 cm thick, lies on a conspicuously bleached layer, 30-50 cm thick. There is a sharp boundary against the underlying indurated pan. This pan is generally double with a brown 2-5 cm thick organic-stained layer over a 10-20 cm thick iron pan. As with most Australian podzols there is no “mor” horizon.
These podzol soils are associated with vegetation of distinctive floristic and structural characteristics. Well over fifty localities containing podzols on the sandstone plateaux around Sydney were then identified, using low slope areas, floristics and vegetation structure as search criteria. The soil type was confirmed from pits and/or auger holes. Two of these sites on the Hornsby Plateau have been chosen for detailed description of the understorey vegetation on the podzols and the type of vegetative changes that occur across a podzol/non-podzol soil boundary. The vegetation characteristics of these podzols have not been reported previously.
SSdy Brooklyn
WX : \ ~~ § \ \
B
Riverview, 9 Ky FCSN
; i New SYDNEY i Guu
|. Wales
Pitt WATER
Main Roads
=) Creeks 7 Salt Water
Warringah Roag
Fig. 1. Locations of the study areas.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 51
SITE DESCRIPTION
The Hornsby Plateau (Fig. 1) is an undulating dissected surface which slopes gently from an elevation of approximately 210 m above Broken Bay in the north to about 120 m in the south about Port Jackson. It consists mostly of the near- horizontally bedded Hawkesbury Sandstone, a sequence of Triassic quartzose sandstones with minor interbedded shale units. It is underlain by rocks of the Narrabeen Group, which outcrop along the north-eastern corner of the Plateau, and is capped in places by remnants of the Wianamatta shale. Meteorological data (Bureau of Meteorology, 1948, 1975) for five stations on or near the Hornsby Plateau are summarized in Table 1.
Site A is in Ku-ring-gai Chase National Park and it is one of twenty-eight podzols in the Lambert Peninsula and McCarrs Creek area. The podzol is in a shallow, basin- shaped catchment. This catchment is located on the plateau surface at 135 m above sea level (a.s.1.) with abrupt rises to 229 m in the north-west and 200 m in the north- east (Figs 1 & 2).
The sand deposit in which the podzol has formed covers approximately 2 ha and is divided by two major creeks (Figs 2b & 3a). The channel of the larger creek has been filled to a depth of about 2 m with layers of iron-rich organic sludge and sand (Fig. 3b). The fill supports a swamp of dense Gahnia steberana and Empodisma minus (syn. Calorophus minor) with other such species such as Banksia robur interspersed.
This sand body is divisible into two units, with the largest section surrounding the northern part of the swamp. This section contains discontinuous podzol development, while the smaller section along the western side of the swamp has a near-continuous, well-developed podzol (Fig. 3a).
TABLE 1
A summary of meteorological data for stations ! on or near the Hornsby Plateau
Rainfall Rain days Daily Max Daily Min (mm) Number Temp Temp Station Yearly total Yearly total (°C) (°C) of means of means Yearly Mean Yearly Mean Marsfield? (33°47'S, 151°7 E) 1178 153 22.4 11.1 45.7ma.s.l. Riverview? (33°50'S, 151°10'E) 928 120 22.9 12.2 22.9maz.s.l. Brooklyn? (33°33'S,151°13'E) 1279 98 — = 5ma.s.l. Hornsby’ (33°42'S,151°06 E) 1237 99 = = 183 ma.s.l. Newport? (83239/S;151°19'E) 1195 113 = = 5ma.ss.l.
1. : The location of these stations is shown on Fig. 1 2. : Bureau of Meteorology (1975) 3. : Bureau of Meteorology (1948)
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
52 VEGETATION ON PODZOLS
From the edge of the sand body to the creek there is a steep slope (approximately 10-18°) and in many places along this slope the podzol pan is revealed at the surface. On the sand body proper the slope is gentle and rarely exceeds 4°. The section across the swamp and podzol (Fig. 3b) shows the sand deposit as a crest 4-5 m high between the swamp and an intermittent drainage line. Further north, in the area where the vegetation was sampled, no change in slope occurs at the sharp boundary between the non-podzolized, yellow-brown sands (non-podzol) and the podzol (Figs 3b & 10). A description of these soil types is given in Appendix 1.
The podzol supports a woodland (Specht, 1970) of Eucalyptus gummifera, Angophora costata and E. piperita, while the non-podzol that surrounds the podzol/swamp complex supports a low woodland to woodland mostly dominated by Eucalyptus haemastoma, E. oblonga, E. gummifera and E. stebert. Ceratopetalum gummiferum is a conspicuous understorey species on the podzol.
Site B is located towards the southern margin of the deeply incised Deep Creek
ES Creeks o Main Roads
yi Dovel Bay Sey Tracks UY, <o (j& sy Salt Water
Heights are in metres
Drawn from Australia |!:25000 series : Broken Bay 9130-1-N, Cowan 9130-IV-N, Mona Vale 9130-!-S and Hornsby 9130-IV-S
Fig. 2. Site A. (a) an enlargement from Fig. 1. (b) an enlargement of (a).
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 53
catchment (Fig. 1). The site (57 ma.s.1.) is within a small basin surrounded by steep, sandstone-benched slopes which rise more than 100 m a.s.]. towards the west. The basin drains eastward into one of the main tributaries of Deep Creek (Fig. 4). A quartz sand deposit up to 4 m deep occurs within the basin and has a general surface slope of 2-3° to the east. The basin is drained by two intermittent streams, of which the more northerly one has trenched into the sand deposit along its northern margin to a depth of 3m. (Fig. 5).
A podzol is developed in the central thicker portion of the sand body and covers 0.15 ha (Fig. 5a). There is an abrupt boundary between the central podzol and the surrounding non-podzolized, yellow-brown sands (Appendix 1).
The podzol supports a woodland of Angophora costata, Eucalyptus gummifera and E. globozdea, while the surrounding non-podzol has a low woodland of E.
Approximate edge of exposed bedrock around the sub-catchment
Drainage line
Podzol and associated vegetation
Discontinuous Podzol and associated vegetation
Non podzolized sands
Vegetation sampling grid (Fig 8)
Transect ( Fig 3b)
Old track to West Head
Swamp
E H
O Coarse sand Swamp fill Horizontal i/ 4 m BRB33 Podzol pan (///\ Bedrock venice Sand & clayey sand Vertical exaggeration 5 X
Fig. 3. Site A. (a) an enlargement from Fig. 2. (b) transect across the podzol and its surroundings.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
54 VEGETATION ON PODZOLS
haemastoma, E. gummifera and E. globoidea. Ceratopetalum gummiferum is again a conspicuous understorey shrub on the podzol.
METHODS FIELD METHODS Sols. The depth of the podzol pan and the lateral extent of the podzol were
determined from trenches, auger holes and by probing with a 7 mm diameter steel rod. The pan could be distinctly felt with the rod. At Site B a detailed examination of
Creek Road Track Saltwater
Drawn from Australia !:25 OOO series : Mona Vale 9130-I-S and Hornsby 9I30-IV-S
Fig. 4. Site B. (a) an enlargement from Fig. 1. (b) an enlargement of (a).
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 55
the soils was carried out and a description of the podzol and non-podzol soils from this site is given in Appendix 1.
Vegetation. Trees were not sampled as they were in insufficient numbers on the well- developed podzol at both sites to give statistically significant results. The shrub layer was sampled using 100 circular quadrats, area 2 m’, set out in a rectangular grid with the longer side at right angles to the abrupt soil boundary. Half the grid and thus half the quadrats were on each soil. The distance between the centres of the quadrats was 3 m in the columns arranged parallel to the longer sides and 2 m in the rows parallel to the soil boundary. At Site A the grid consisted of five columns and twenty rows. At Site B it consisted of seven columns and fourteen rows, with an extra quadrat added to the
GM SC«Vegetation sampling Approx. edge of aie ( fig.9 ) exposed bedrock E
Transect (Fig. 5b) Drainage line
Podzol and associated vegetation
Non podzolized sands
O 50m —
Coarse sand
Loamy coarse sand Loamy sand : pan of podzol Sandy loam
Clayey sand
Sandy clay
Sandstone bedrock
Sample points expressed in terms of principal prolific
=< Gn22!
Fig. 5. Site B. (a) an enlargement from Fig. 4 b. (b) transect across the podzol and its surroundings.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
56 VEGETATION ON PODZOLS
podzol and non-podzol to give a total of 100 quadrats. The variation in the grid pattern was dictated by the size and shape of the podzols. Row numbers are shown on Figs 10 & 11.
The small quadrat size was chosen to facilitate sampling of the small understorey species. All dicotyledon and gymnosperm stems rooted in the quadrats and with a diameter of 4-100 mm 20 cm from the ground, were measured to the nearest millimetre. The height of each stem was estimated to the nearest 0.5 m. The monocotyledons Xanthorrhoea arborea and X. medza were only recorded as present at Site A, but at Site B the number of individuals of this genus was recorded also.
TREATMENT OF DATA
In order to reduce some of the minor variation in the data, species occurring in less than 5% of the total number of quadrats at each site were excluded from the relative importance calculations and the association and principal component analyses.
Relative Importance and Exclusiveness. The frequency, density (Kershaw, 1973), average basal area and average height were calculated for each species on each soil at both sites (Tables 2 & 3). The relative importance value (RIV) for each species was obtained by summing the relative density, relative frequency, relative dominance (Kershaw, 1973), and the relative height to give a total out of 400. The relative height was calculated by:
height of sp. on a particular soil x 100 2 height of all spp. on a particular soil
The RIV is similar to the “importance value” used by Curtis and McIntosh (1950). Relative height was added to give a better balance between size (dominance and height) and numbers (frequency and density) .
The exclusiveness value is defined as:
number of quadrats in which species A occurred on a particular soil number of quadrats in which species A occurred
and was calculated for the soil on which the species was most frequent. It is a measure of the degree to which a species is restricted to a particular soil in the sampled area. Species with an exclusiveness value of greater than 85% were assigned to the soil where they were most frequent.
All species mentioned are listed in Appendix 2.
Principal Component Analysis. Each quadrat was weighted by RIVs obtained by summing the relative density, relative dominance and the relative height of each species. The resulting RIV data for the spp. within the quadrats were subjected to principal component analysis, using a programme based on the BMDOIM programme of Dixon (1973). The small quadrat size ensured maximum variability (Table 4) and the plot of these results is not included. The quadrats were then grouped into rows parallel to the soil boundary, the RIV, also including relative frequency, was calculated for each species, and the data were then re-analysed (Table D))
Association Analysis. The normal and inverse association analyses were applied to the data using the DIVINFRE program of the CSIRO Division of Land Research, Canberra (see Williams, 1976, p.94). This is a divisive monothetic analysis based on presence/absence data. The program was terminated at the five group level, as it was more than sufficient to show the difference in the vegetation on the podzol and non- podzol. The inverse analysis added little to the information included in Tables 2 & 3
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN AND G.S.HUMPHREYS
Species
TABLE 2
Frequency, density, average basal area, average height, relative importance value (RIV) and exclusiveness of species on the podzol (P) and non-podzpol (N-P) at Site A.
Av.basal Av. height
(2 m) P N-P
P
area (cm?) N-
RI (40 P
57
{Xanthorrhoea arborea
*Pimelea linifolia *Angophora costata *Eucalyptus piperita Ceratopetalum gummiferum Gompholobium latifolium - Xylomelum pyriforme Phyllota phylicozdes Ricinocarpos prnifolvus Acacia suaveolens A. terminalis A. ulicifolia
i)
Podzol species
Boronia pinnata
Hibbertia linearis Leucopogon ericozdes Dillwynza retorta Platysace linifolia Banksza serrata +Xanthorrhoea media
Common species
Leptospermum attenuatum p : é :
Lambertia formosa Petrophile pulchella Epacris pulchella Leucopogon microphyllus Banksia spinulosa Grevillea sericea Pultenaea elliptica Gompholobium grandiflorum Epacridaceae sp. Acacia myrtifolta Grevillea buxifolia Eucalyptus gummifera
*Boronza ledzfolia
*Hakea gibbosa
*H. dactylozdes
*Eucalyptus sp.
*Leptospermum flavescens
*Woollsia pungens
*Lomatia silatfolia
*Persoonza levis
*P. lanceolata
*Banksza asplenzfolia
*Fucalyptus haemastoma
*Leptospermum squarrosum
*Persoonia sp.
*Bossiaea heterophylla
{Overall average
Frequency (%) 1p INGLY 30 0 4 0 2 0 2 0 66 0 16 0 10 0 42 2 42 4 34 4 18 2 16 2 22 8 360.22 340s 28 20 820 6 8 26 =. 30 30 = 40 22) 950 2 16 = AKY 0 40 0 32 0 24 0 20 0. 18 0 16 0 12 0 8610 0 10 0 8 10 0 8 0 8 0 8 0 8 0 6 0 6 0 6 0 6 0 4 0 4 0 4 0 2 0 2 0 2
Density No/100m? P N-P — 0 sO 1 oO 1 0 184 0 2 nO 14 O YB & 118 2 ye 10 1 135 71 ay @) 7a) © 28) 61 38 eels 8) lil 31 23 48 108 5 54 1 61 0 32 0 30 0 34 0 15 0 31 0 21 0 14 0 9 0 9 0 5 0 4 0 4 0 24 OR 0 4 0 3 0 8 OQ §& On 3 0 8 O°. 2 @ 1 Opell 0 1
+Monocotyledons: not included in RIV calculations *Species that occurred in less than 5% of all the quadrats: not included in RIV calculations {The overall average does not include species marked*
— ©
bo bo
— ~JI
oo
—" oO
_ So
Non-Podzol species
Vv 0) Exclu- N-P siveness — 100 — 100 — 100 — 100 0 100 0 100 0 100 2 95 2 91 2 89 1 90 1 99 5 73 14 62 20 55 10 50 6 57 56 54 = 57 68 69 27 89 45 96 20 100 17. —-:100 25 100 12 100 18 100 11 100 7 100 6 100 9 100 15 100 — 100 — 100 — 100 — 100 = § iO) — 100 = ia = IO =) 00 = 100) = io = i100 = 100) = N00
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
58 VEGETATION ON PODZOLS
TABLE 3
Frequency, density, average basal area, average height, relative importance value (RIV) and exclusiveness of species on the podzol (P) and non-podzol (N-P) at Site B.
Frequency Density Av.basal _ Av. height RIV
(%) No/100m? area (cm?) (% m) (400) Exclu- Species Peo NePi Bo NeP Ps N-P. P N-P P_ N-P siveness {+Xanthorrhoea arborea 24 0 13 — 100° +X. media 16 0 So 100 *Dodonaea triquetra 8 0 10 +0 0.7 0 2.0 0 — — 100 < *Grevillea linearifolia 6 0 0 1.3 0 2.5 0 — = 100 & *Xylomelum pyriforme 6 0 Bb oO 1.6 0 1.0 0 - — 100 = Podocarpus spinulosus 42 0 41 0 0.2 0 0.5 0 21 NOD Ceratopetalum gummiferum 30 0 35 0 8.0 0 3.5 0 55 QO 100 € Gompholobium latifolium 16 0 330 0.6 0 2.0 0 24 0 100 Ricinocar pos pinifolcus 10 0 42 0 0.3 0 1.5 0 24 0 100 *Leucopogon ericoides 4 2 11 3 0.2 0.2 1.0 1.0 — — 67 *Angophora costata 4 2 2 1 oo) Boh “SO 40 - _ 67 Woollsia pungens 22 16 18 10 ORS Os4 270220 14 9 58 Banksta serrata 16 14 Ik 32 IO GO Me We BO NE ae). Dillwynia retorta 72 64 145 76 09) OO. Ao Ao 947-9245) 53 +e Hibbertza sp. 8 8 Abe Si 0.5 0.5 Waly Le) 7 4 50 &. Ertostemon australastus 18 18 15 11 0.8 O49 Ao 1.8 16 8 i) Petrophile pulchella 22 24 14 21 Boil oN 3.0. 2.5 16 9 52 g Acacia suaveolens 18 20 12 #14 0.4 1.0 1.0 2.5 14 12 53 §€ Grevillea buxtfolia 4 6 B8) & 4:0 1.0 3.5 2.5 3 5 60 6 Eucalyptus gummifera 6 10 6 Uo 28st BO 40 2B 63 E. sp. 2 4 We Bo BBO 9.9 BO) Bs — - 67 Acacia terminalis 4 8 2 4 9.7 12.1 4.5 4.4 6 16 67 Leptospermum attenuatum 28 62 22 DD 5.8 8.9 3.0 2.5 32 73 69 Banksia marginata 16 96 11 127 0.8 0.7 2.5 2.0 11 68 86 Phyllota phylicotdes 4 28 4 46 0.3 0.4 1.5 1.0 3 23 88 Banksia ertcifolia 2 18 LS e213 86) a 4053.0) Anyi 90 Hakea gibbosa Bh Ah 3 23 38 1259 3.5 3.0 Cre 2 89 Platysace linearifolia 0 20 0 20 0 0.4 #0 2.0 0 14 100 § Lambertia formosa 0 12 0 26 0 | Oe) 1.0 0 13 100 > Grevillea sericea 0 12 0 6 0 05 O 2.0 0 LOO R Hakea teretifolia 0 10 0 5 0 (aos) AY) 3.5 0 8 100 8 *Pultenaea sp. 0 4 0) 2 0 0.1 O 2.0 — — 100 & *Hakeoa dactyloides 0 4 0 2 0 2.6 0 2.4 = = 100 *Kunzea capitata 0 2 0 1 0 0.5 0 3.5 — 00S *Leptospermum sp. 0 2 0 1 0 0.3 0 2.0 = — 100 *Conospermum longifolium 0 2 0 en wo) 06 O 2.0 — — 100 *Pultenaea elliptica 0 4 Ona 0 06 O 2.5 — — 100 {Overall average ZelB) oe) Za) Pao)
+Monocotyledons: not included in RIV calculations *Species that occurred in less than 5% of all the quadrats: not included in RIV calculations {The overall average does not include species marked*
and it is not presented in the results. All computations were run on a UNIVAC 1100 computer at Macquarie University. RESULTS TABLES 2 AND 3 When species with an exclusiveness value of greater than 85% were assigned to the soil where they were most frequent, three distinct groups which were readily visible in the field resulted: podzol species, species common to both soil types, and non-
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 59
podzol species. This classification applies only to the sampled areas, as none of the podzol species is restricted regionally to this soil.
The ratio of these three groups (podzol: common: non-podzol) for the full species list at Site A is 1.5: 1: 3.25. Not only is the group of species common to both soils the smallest, but it also contributes only a quarter to the total RIV on the podzol and slightly less than half on the non-podzol. At the more sheltered, moister Site B the podzol group is the smallest, with the common and non-podzol groups the same size (1: 1.5: 1.5). At this site the group of species present on both soils contributes well over half to the RIV total on the podzol and slightly over a half on the non-podzol.
Vegetation on the Podzol. The species locally restricted to the podzol and present at each site are Ceratopetalum gummiferum, Ricinocarpos pinifolius, Gompholobium latefolium, Xylomelum pyriforme and Xanthorrhoea arborea.
At Site A, C. gummiferum is the principal species on the podzol. This single species contributes 70% of the summed values for the basal area, 40% for height, 26% for density and 16% for frequency to give a total score of 152 out of 400. The next two most important plants on this soil, R. penzfolius (RIV 42) and Phyllota phylicoides (RIV 35), attain a score of less than a third of that for C. gummzferum. Although these two podzol species are quite common, their small size makes them a less important and less conspicuous part of the flora.
At Site B there is no clear principal species on the podzol. A species common to both soils, Dzllwynza retorta, has a score of only 94, with C. gummiferum reaching a value of over half this score at 55. Another species common to both soils, Leptospermum attenuatum, is the third most important species (RIV 32). The regionally less-common species Podocarpus spinulosus is confined to the podzol at Site B where it is the second most frequent species.
C. gummiferum is less important at the more sheltered Site B than at Site A even though it is usually found in moist, sheltered places. At Site B the plants were larger (average height and basal area 3.5 m and 8 cm’ respectively) than at Site A (average height and basal area 2.5 m and 6 cm’) but only 35 stems were recorded at Site B compared with 184 stems at Site A. This difference may be caused by a lower fire
TABLE 4
Principal component analysis, in quadrats (100) of vegetation RIV data: table of components and their respective eigenvalues
site component eigenvalue % variance cumulative variance accounted for % A 1 2.5169 9 9 2 1.9165 7 16 3 1.6483 7 23 4 1.4765 5 28 5 1.4134 5 33 6 1.3181 5 38 7 1.2825 5 43 8 1.1935 4 47 B 1 2.2376 10 10 2 1.6529 7 17 3 1.5981 7 24 4 1.5219 6 30 5 1.4715 7 37 6 1.3452 6 43 7 1.1847 5 48 8 1.1707 5 53
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
60 VEGETATION ON PODZOLS
frequency, as plants in fire-prone areas tend to have larger numbers of smaller shoots than plants in less fire-prone areas.
P. phylicozdes is the only species that grows on the opposite soil at each site but a few species are confined to one soil at one site and common to both soils at the other.
Vegetation on the Non-podzol. A species common to both soils, L. attenuatum, is the most important species on the non-podzol at both sites.
At Site A there is no predominant species on the non-podzol. L. attenuatum and another species common to both soils, Banksza serrata, are the most important species with RIV totals of 68 and 56 respectively, while the non-podzol species, Petrophile pulchella, has a score of 45. There are a large number of infrequent non-podzol species at this site.
At Site B, L. attenuatum has a RIV of 73. The next most important species, Banksia marginata, is mostly restricted to this soil and has a score of 68, while another species common to both soils, Dillwynza retorta, has a RIV of 45. B. marginata has a very high frequency (98%) and is therefore a conspicuous species on this soil even though its RIV is not large.
PRINCIPAL COMPONENT ANALYSIS
The first component of the row analysis accounted for over 30% of the variability and with the second component over 40% at both sites (Table 5) with the first component representing the change in soil type.
The Site A analysis shows three groups (Fig. 6), the podzol group and two groups on the non-podzol. The two podzol border rows, nine and ten, are included within the podzol group but the non-podzol border rows, eleven, twelve and thirteen, lie midway between the podzol group and the other non-podzol rows along the first component. In these three rows, species common to both soils are important ingredients and many of the non-podzol species are absent. Within the non-podzol group there is a gradation from rows fourteen to twenty along the second component. This may relate to drainage and slope.
At Site B the podzol group is split into two sub-groups, with the two rows on the steeper slope adjacent to the creek well separated from the other podzol rows along the
TABLE 5
Principal components analysis, in rows, of vegetation RIV data: table of components and their respective eigenvalues
site component eigenvalue % variance cumulative variance accounted for % A 1 8.5464 32 32 2 3.0693 11 43 3 2.5654 10 53 4 2.3691 8 61 5 1.7961 7 68 6 1.5891 6 74 7 1.5119 5 79 8 1.3026 5 84 B 1 7.0695 31 31 2 3.1869 14 45 3 2.9750 13 58 4 2.2209 9 67 5 1.8840 8 75 6 1.7358 8 83 7 1.4994 6 89 8 0.6869 3 92
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN AND G.S. HUMPHREYS 61
Fig. 6. Site A. Plot of the lst x 2nd principal component scores of the twenty rows. The location of rows is shown on Fig. 10. © Podzol row A Non-podzol row.
Fig. 7. Site B. Plot of the Ist x 2nd principal component scores of the fourteen rows. The location of rows is shown on Fig. 11. © Podzol row A Non-podzol row.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
62 VEGETATION ON PODZOLS
second component (Fig. 7). In both rows, two out of the three most important species are common to both soils but these rows contain the only intrusion of a non-podzol species, Hakea gibbosa. Podzol species such as C. gummiferum and R. pinifolius are also present and the mixture of podzol and non-podzol species guaranteed that these two rows would be plotted well away from the main groups.
The podzol border rows, six and seven, are within the main podzol group but the two non-podzol border rows, eight and nine, lie between the podzol and non-podzol groups along the first component. As for Site A, the non-podzol border rows are dominated by species common to both soils, while many non-podzol species are absent. Although the podzol and non-podzol plot out as two distinct groups there is a rough ordering from rows three to fourteen along the first component.
NORMAL ASSOCIATION ANALYSIS
The first two divisions in the normal association analysis at Site A define the podzol vegetation, while at Site B the first division basically divides the vegetation into podzol and non-podzol (Figs 8 & 9).
The community maps (Figs 10 and 11) show that only 6% of the quadrats at Site A and 10% of the quadrats at Site B do not conform to the soil groups. Even this small amount of non-conformity seems to be the result of the highly artificial classification system, as the chance occurrence of one species in a quadrat can dictate its final grouping. When the relative importance values of the species are considered most of
LEGEND
+ Positively associated - Negatively associated N No. of quadrats
Ceratopetalum gummi ferum
= 2 uJ = 2 [e) O 2 < E a = S re =
Epacris itt if Uf Ricinocarpos bay ate) Pinifolius Hibbertia linearis
Se anoe Sooes
Mapping group No. as used on Fig. !0 b.
Fig. 8. Site A. Normal association analysis.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 63
these apparent discrepancies disappear. Even the well-defined finger of non-podzol vegetation that intrudes into the podzol on the Site B community map is the intrusion of a single species, B. marginata. This species is unimportant in these quadrats.
DISCUSSION
All results, whether those of the exclusiveness calculations, the association analysis or the principal components analysis, showed that there was a difference in the vegetation on the podzol and the non-podzol soils at each site. This difference was less evident at the moister, more sheltered Site B than at the plateau site, Site A. At Site B there were more species common to both podzol and non-podzol soils, and these species were more important than at Site A. The vegetation characteristic of the podzol soil is present to the podzol boundary where it ceases abruptly. At both sites, on the non-podzol soil the vegetation close to the soil junction is dominated by species common to both soils, with many of the non-podzol species being infrequent. Though the two podzol sites differ in topographic terms, the vegetation on both podzols looks remarkably similar. Extensive field reconnaissance within the Sydney Basin has shown that the vegetation on these two sites is typical of most podzols on the sandstone plateaux.
LEGEND
+ Positively associated — Negatively associated N No. of quadrats
Banksia marginata
Dillwynia retorta
pe 2 ud = 2 1e) O 2 2 Ee <x = S u_ =
Podocarpus spinulosus
Mapping group No. as used on Fig I! b
8 ese- sold
@Q cesossoea IS) Geocesce
Fig. 9. Site B. Normal association analysis.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
64 VEGETATION ON PODZOLS
D oa D C
MAPPING UNIT MAPPING GROUP : as in fig 8
(ee Podzo! Mostly associated with the podzol soil.
Mostly associated with the
Non-podzolized sands non-podzol soil
Soil boundar _ First division level of the
normal association analysis
[OM ‘Contour interval O:25m Arbitary datum point at A is approximately !35m. a.s.l
Fig. 10. Site A. (a) Soil and microtopography of the sample grid (Fig. 3a) (b) Vegetation community map.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S. HUMPHREYS 65
The podzol vegetation is usually woodland or open-forest and is often taller than the trees on the surrounding soils. The tree species common to both sites, Angophora costata and Eucalyptus gummifera, are usually important components on sandstone podzols over most of the Sydney region. The third species, although frequently Eucalyptus piperita, is less constant. The understorey species restricted to the two podzols, Ceratopetalum gummiferum, Xylomelum pyriforme, Xanthorrhoea arborea and to a lesser extent Rzcinocarpos pinifolius and Gompholobium latifolium, are typical podzol species. Banksza serrata is often an obvious species on podzols and was present at both sites. Other species may be locally important.
The important species on these podzols are usually plants with a wide habitat range, or are those which occur on the upper and middle gully slopes. C. gummzferum thrives in the damper gullies (Rotherham e¢ al., 1975) but it is also found in three situations on and near the plateaux surface. The first is the rocky positions, either along creeks incised a few metres below the immediate plateau surroundings, or on the sheltered sides of rock outcrops. The second situation is the deep sand bodies, most of which have podzol soils, where rock outcrops are noticeably absent. Thirdly, C.
MAPPING UNIT MAPPING GROUP : as in fig 9
Fxg Podzol Mostly associated with the podzol soil.
ostl iated with the Non-podzolized sands Bo eat ih
ig colle boundany First division level of the
normal association analysis
10 m Contour interval O-25m
Arbitary datum point at A is approximately 57 m a.s.!.
Fig. 11. Site B. (a) Soil and microtopography of the sample grid (Fig. 5a) (b) Vegetation community map.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
66 VEGETATION ON PODZOLS
gummiferum is sometimes found on more clayey soils, either derived from occasional intercalations of shale, or associated with the boundary between Hawkesbury Sandstone and the overlying Wianamatta shale, such as along Mona Vale Road between the junctions with Forest Way and McCarrs Creek Road (Fig. 1).
On the Hornsby Plateau X. arborea and G. latifolium are most frequent on upper and middle slopes, while the very common B. serrata has a wide habitat range, as do the less common X. pyriforme and R. pinifolius. Other common species on podzols include the habitat-tolerant Leptospermum attenuatum and Dillwynza retorta and the slope species Smilax glyciphylla and Platylobium formosum.
This assemblage of species may also occasionally be found on sand deposits some of which may have a considerable clay component but no development of a podzol profile. In general, however, a combination of topographic situation (low slope) and floristics may be used to locate podzols.
Grevillea species have rarely been recorded on the main body of a podzol. The absence of this genus is one of the most unexpected features, as twenty species with a wide range of habitats occur in the Sydney region (Beadle, Evans and Carolin, 1972). One of these species, Grevillea lineartfolza, is a very common plant of the gully slopes on the Hornsby Plateau. In this situation it frequently grows with C. gummiferum. No Grevillea species were recorded on the podzol at Site A, although two species grew on the adjacent soil. At the Site B podzol, G. lineartfolia and G. buxzfolia grew only on the steeper slope adjacent to the creek.
Many species common in the heaths and low woodland on the plateau are infrequent on podzols, even though the podzols may immediately adjoin such communities. The very common plateau tree, Eucalyptus haemastoma, has rarely been recorded, while shrub species uncommon on the podzol include Banksia asplenifolia, Gompholobtum grandiflorum, Angophora hispida and the Grevillea species.
Because the podzol vegetation has greatest affinity with the gully slope vegetation and very little with the plateau vegetation, podzols on the plateau are easier to locate than those in the gullies. In plateau areas the extent of the podzol can be roughly mapped by the distribution of the easily observed C. gummizferum which is usually the most conspicuous species on these podzols even where it is not common. It is a tall slender plant (average heights of 2.5 m and 3.5 m at Site A and Site B) and stems up to 4 m tall are present on both podzols. On the podzols, C. gummiferum is usually multi-stemmed and the dense, light green leaves are often present from near ground level. This tall dense mass contrasts sharply with the sparser, grey-green xeromorphic foliage of most species. Although the average height of vegetation on the podzol and non-podzol at each site was similar (1.5 m and 1.0 m at Site A and 2.5 m at Site B; Tables 2 and 3), the presence of this tall dense mass of C. gummiferum foliage usually makes the vegetation on the podzols appear taller than the surrounding vegetation.
On the podzols R. pinzfolius and G. latifoltum are fairly inconspicuous, thin- stemmed shrubs rarely more than 2 m tall. B. serrata may be a small tree up to 4-5 m tall but the average height of this thick-stemmed species was only 1-1.5 m. X. arborea is a fairly obvious member, as individuals with stems up to 2 m tall are often present. Xylomelum pyriforme is usually present but is always uncommon.
The podzol vegetation is a fairly stable unit, as all of the six indicator species regenerate from parent plants after fire. C. gummiferum often has a clumped distribution as it usually possesses a well-developed lignotuber, especially in fire-prone areas such as the plateau surface. The lignotuber of one plant may be almost as large as the 2 m? quadrat used. Small B. serrata and X. pyriforme regenerate from lignotubers, but larger plants usually regenerate from epicormic buds on the main
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 67
stem. R. pinifolius and perhaps G. latifoltum regenerate from underground perennating buds and are clumped on a much smaller scale than C. gummiferum. X. arborea continues growing from its apical meristem and all the trees regenerate from lignotubers or epicormic buds after fire.
This paper unlike that of Enright (1978) does not deal with important general issues such as the genesis of podzol soils, the origin of the characteristic vegetation or the interaction between vegetation and soils. Nevertheless we would like to consider briefly two possible causes of the different vegetation between podzols and non-podzol soils. The different distribution of plants on soils with essential chemical differences is well documented. Thus in the field situation dealt with by Enright (1978) both calcareous and non-calcareous parent materials occurred, with podzols, and hence podzol vegetation, best developed on the non-calcareous aeolian sands. Such a dramatic difference in soil chemistry is not likely at Sites A and B as the acidic parent material of the podzol and non-podzol is all derived from the Hawkesbury Sandstone. However, at Sites A and B there is a difference in soil drainage characteristics between the podzol and the non-podzol. The podzol has a well-aerated and well-drained layer overlying a less pervious pan that may also act as a perched water table. In comparison, the non-podzol is a less well drained soil. This contrast may explain some of the differences in the vegetation between the podzol and the non-podzol.
ACKNOWLEDGEMENTS
Our thanks are due to Dr. M. Westoby for criticizing an early draft of this manuscript and Dr. D. A. Adamson, School of Biological Sciences, Macquarie University; to Mr. T. R. Paton, Mr. P. B. Mitchell and the cartographic staff, School of Earth Sciences, Macquarie University; Dr. R. Outhred, Institute of Technology, N.S.W.; and the Royal Botanic Gardens of N.S.W.
References
BEADLE, N. C. W., Evans, O. D., and Caron, R. C., 1972. — Flora of the Sydney Region. Sydney: A. H. & A. W. Reed.
BUCHANAN, R. A., 1980. — The Lambert Peninsula, Ku-ring-gai Chase National Park. Physiography and the distribution of podzols, shrublands_and swamps, with details of the swamp vegetation and sediments. Proc. Linn. Soc. N.S.W. 104: 73-94.
BuREAU OF METEOROLOGY, 1948. — Results of Raznfall Observations made in N.S.W. Section I-VI, Commonwealth of Australia. ——, 1975. — Climatic Averages Australia Metric Edition, Dept. Science and Consumer Affairs.
Canberra. Australian Government Publishing Service.
Burces, A., and Drover, D. P., 1953. — The rate of podzol development in the sands of the Woy Woy district, New South Wales. Aust. J. Bot. 1: 4-94.
BurRROuUGH, P. A., BRown, L., and Morris, E. C., 1977. — Variations in vegetation and soil patterns across the Hawkesbury Sandstone plateau from Barren Grounds to Fitzroy Falls, New South Wales. Aust. J. Ecol. 2: 137-159.
Curtis, J. T., and McInrosu, R. P., 1950. — The inter-relations of certain analytic and synthetic phytosociological characters. Ecology 31: 434-455.
Dixon, W.J., (ed.), 1973. — BMD Biomedical Computer Programs. Los Angeles: University of California Press.
ENRIGHT, N. J., 1978. — The interrelationships between plant species distribution and properties of soils undergoing podzolization in a coastal area of S.W. Australia. Aust. J. Ecol. 3: 389-401.
Forster, G. R., CAMPBELL, D., BENSON, D., and Moore, R. M., 1977. — Vegetation and soils of the western region of Sydney, pp. 53. CSIRO Division of Land Use Research. Tech. Memo 77/10. Canberra.
HamILTON, G. J., 1976. — Soil resources of the Hawkesbury River Catchment, New South Wales. J. Sozl Conserv. Serv. N.S.W. 22:L 204-229.
KersHAw, K. A., 1973. — Quantitative and Dynamic Plant Ecology. London: Edward Arnold.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
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NorTHcoteE, K. H., 1974. — A factual Key for the Recognition of Australian soils. 4th ed. Glenside, South Australia: Rellim Technical Publications.
PATON, T.R., 1978. — The Formation of Soil Material. London: George Allen & Unwin Ltd.
PIDGEON, I. M., 1937. — The ecology of the central coastal area of New South Wales. I. The environment and general features of the vegetation. Proc. Linn. Soc. N.S.W. 62: 315-340.
——, 1938. — The ecology of the central coastal area of New South Wales. II. Plant succession on the Hawkesbury Sandstone. Proc. Linn. Soc. N.S.W. 63: 1-26.
——, 1941. — The ecology of the central coastal area of New South Wales. IV. Forest types on soils from the Hawkesbury Sandstone and Wianamatta Shale. Proc. Linn. Soc. N.S.W. 66: 113-137.
ROTHERHAM, E. R., BLAXELL, D. F., Briccs, B. G., and CarRo.in, R. C., 1975. — Flowers and Plants of New South Wales and Southern Queensland. Sydney: A.H. & A. W. Reed.
SPECHT, R., 1970. — Vegetation. In G. W. Leeper (ed.), The Australian Environment. 4th ed. Melbourne: CSIRO & Melbourne University Press.
Stace, H. C. T., Hupsie, G. D., BREWER, R., NoRTHCOTE, K. H., SLEEMAN, J. R., MuLcany, M. J., and HALLSwortH, E. G., 1968. — A Handbook of Australian Soils. Glenside, South Australia: Rellim Technical Publications.
WALKER, P. H., 1960. — A soil survey of the County of Cumberland, Sydney Region, New South Wales.
Soil Surv. Unit Dept. Agric. N.S.W., Bull. 2: 7-109. , and Hawkins, C. A., 1957. — A study of river terraces and soil development on the Nepean River, N.S.W./J. Proc. R. Soc. N.S.W. 91: 67-84.
WILLIAMS, W. T., 1976. — Hierarchical divisive strategies. In W. T. Williams (ed.), Pattern Analysts in
Agricultural Science. Melbourne: CSIRO & Elsevier Scientific Publishing Company.
APPENDIX I
The soils were mapped as layers and described in terms of field texture, matrix fabric, planar void development, consistency (as defined in Paton, 1978), Munsell colours, pH and nature of the boundaries (as defined in Northcote, 1974). Two “Type Profiles” (Paton, 1978), one of a podzol and the other of a non-podzol, are described from the vegetation sample area at Site B. These soils were also classified into “Great Soil Groups” (Stace et al., 1968) and “Principal Profile Forms” (Northcote, 1974).
There is a sharp boundary of less than 2 m between these two soil types and the intravariation is small.
Type Profile : Podzol Great Soil Group: Podzol Principal Profile Form: Uc 2.3, approx. Uc 2.36
Location: Site B; centre of the podzol half of the sampling grid (Figs. 5 & 11). Parent Material : One to 2 m of sand deposits with some cobbles overlying 2 m of zn sztu? altered sandstone. Vegetation: Shrubs of Ceratopetalum gummiferum, Dillwynia retorta, Xanthorrhoea _ media and a nearby dead stump of Eucalyptus gummifera? and other typical podzol vegetation.
Description:
2-0 cm Leaf litter of the above-mentioned species, especially Ceratopetalum
(leaf litter) gummiferum and Xanthorrhoea media; white fungi hyphae connecting many leaves; sharp even to:
0-21 cm Organic-rich coarse sand (silky feel); 2.5Y2/1 (dry and moist) ; single
pH 5.0-5.5 grain sand with minor coherence from the dense root mat; no planar voids ; apedal and fragile consistency; abundant charcoal; earthworms in the upper 10 cm, white Scarabaeidae grub in the lower, diffuse wavy to:
21-64 cm Coarse sand; 10YR7/1 (dry), 6/2 (moist) ; incoherent single grain sand ;
(bleached layer) no planar voids; apedal and very fragile consistency; roots are sparsely
pH 6.0-6.5 distributed and are mainly confined to the upper 15 cm; small patches of fungi hyphae linking individual sand grains in lower 10-20 cm; sharp wavy to: j
64-70 cm Coarse loamy sand; 10YR3/4 (dry), 4/3 (moist) ; mottled with 10YR2/3
(organic pan)
(dry and moist) ; matrix fabric, weakly coherent, non-uniform and highly porous; planar voids are absent; consistency, apedal and brittle when dry; small patches of fungi hyphae binding individual grains; within and beneath a decaying root (5-7 cm diameter) it is 1OYR2/3; sharp irregular to:
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
70-89 cm (iron pan) pH 6.0
89-100 cm pH 5.5
Type Profile :
Great Soil Group: Principal Profile Form: Location:
Parent Materzal : Vegetation: Description :
1.0 to0cm
0-8 cm pH 5.0
21-50 cm pH 5.0
50-180 cm pH 5.5-6.0
GYMOSPERMAE Podocarpaceae :
R. A. BUCHANAN AND G.S.HUMPHREYS 69
Coarse loamy sand; 10YR 4/4 (dry), 3/4 (moist) with mottles of 10YR 5/6 (dry), 4/6 (moist); matrix fabric, moderately coherent, non- uniform and very porous; no planar voids; consistency, apedal and brittle when dry. Interspersed are mottles of 7.5 YR 5/8 (dry & moist) ; matrix fabric, very coherent non-uniform and porous with no planar voids in the upper part of the layer but it is very nodular in the lower half; consistency, brittle and very indurated when dry requiring considerable pressure to disrupt. Charcoal throughout often centering the very indurated material; earthworms and termites present :
Coarse clayey sand; 10YR 5/8 (dry & moist); matrix fabric, mod- erately coherent, non-uniform; no planar voids; consistency, slightly plastic; some charcoal and few roots.
Depositional Earth
Earthy Sand
Gn 2.21 to Um 5.22
Site B, near centre of the non-podzol half of the sampling grid (Figs 5 & iL).
Approximately 0.5 m of sand deposits overlying more than 1 m of zn sztu? altered bedrock.
Small tree of Persoonza levis and shrubs of Banksta marginata, Dillwynia retorta, Leptospermum attenuatum and Platysace lineartfolia, and other typical non-podzol vegetation.
Thin litter layer, mainly of leaves from Persoonza levis; sharp even to: Coarse sand; 2.5Y 4/2 (dry), 5/2 (moist); highly porous, sand- dominated, non-uniform, fragile matrix fabric, with added but weak coherency due to the organic matter; no planar voids; consistency apedal and very brittle; roots throughout; plenty of coarse charcoal; grub (Scarabaeidae) present; clear, even to:
Medium sandy loam; 10YR 5/5 (dry and moist); porous, sand- dominated, non-uniform and coherent matrix fabric with indurated nodules 10YR 4/6 (dry and moist) of weakly porous to dense, sand- dominated, non-uniform and very coherent matrix fabric; no planar voids; consistency apedal and brittle; fine charcoal; some white quartz pebbles one to 1.5 cm diameter; few large roots; gradual even to:
Clayey sand 10YR 5/8 (dry and moist); moderately porous clay- dominated, non-uniform and coherent matrix fabric; no planar voids; slightly subplastic consistency; occasional white quartz pebble one to 1.5 cm diameter. In lower 1 m (augered) mottled streaks of 7.5 YR 5/8 dry and moist occur.
APPENDIX II
Species mentioned in paper
Podocarpus spinulosus (Sm.) R. Br. ex Mirb.
ANGIOSPERMAE Dicotyledones Proteaceae:
Banksza aspleniifolia Salisb.
B. ericifolia L.£.
. robur Cav. . serrata L.f. . Sptnulosa Sm.
maaowh
. marginata Cav.
Conospermum longifolium Sm. ssp. longifolium
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
70 VEGETATION ON PODZOLS
Grevillea buxifolia (Sm.) R. Br.
G. lineartfolia (Cav.) Druce
G. sericea (Sm.) R. Br.
Hakea dactyloides (Gaertn.) Cav.
H. gibbosa Cav.
H. teretzfolia (Salisb.) J. Britt.
Lambertia formosa Sm.
Lomatza stlazfolia (Sm.) R. Br.
Persoonza lanceolata Andr.
P. levis (Cav.) Domin
P. Unnamed
Petrophile pulchella (Salisb.) Knight
Xylomelum pyriforme Sm. Dilleniaceae:
Hibbertza linearis DC. var. obtuszfolia
H. sp. Euphorbiaceae:
Ricinocarpos pinifolius Desf. Cunoniaceae:
Ceratopetalum gummiferum Sm. Mimosaceae:
Acacia terminalis (Salisb.) MacBride
A. myrtifolia (Sm.) Willd.
A. suaveolens (Sm.) Willd.
A. ulicifolia (Salisb.) Court Papilionaceae:
Bosstaea heterophylla Vent.
Dillwynza retorta (Wendl.) Druce var. retorta
Gompholobium grandiflorum Sm.
G. latefolium Sm.
Phyllota phylicoides (Sieb. ex DC.) Benth.
Platylobtum formosum Sm.
Pultenaea elliptica Sm.
P. st¢pularis Sm. Myrtaceae:
Angophora hispida (Sm.) Blaxell
A. costata (Gaertn.) Druce
Eucalyptus globoidea Blakely
E. gummifera (Gaertn.) Hochr.
E. haemastoma Sm.
E. oblonga DC
E. piperita Sm. ssp. piperita
E. steber L. Johnson
E. sp.
Kunzea capitata Reichb.
Leptospermum attenuatum Sm.
L. flavescens Sm.
L. squarrosum Sol. ex Gaertn. Rutaceae:
Eriostemon australaszus Pers.
Boronza ledifolia (Vent.) J. Gay
B. pinnata Sm. Sapindaceae:
Dodonaea triquetra Wendl. Umbelliferae :
Platysace lineartfolia (Cav.) C. Norman Epacridaceae:
Epacris pulchella Cav.
Epacridaceae sp.
Leucopogon ericozdes (Sm.) R. Br.
L. microphyllus R. Br.
Woollsia pungens (Cav.) F. Muell.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R.A. BUCHANAN AND G.S.HUMPHREYS 71
Monocotyledones
Xanthorrhoeaceae : Xanthorrhoea arborea R. Br. X. media R. Br. ssp media
Cyperaceae: Gahnia steberana Kunth
Restionaceae : Empodisma minus (Hook.f) Johnson & Cutler syn. Calorophus minor Hook.f
The species names are those currently used by the Royal Botanic Gardens of N.S.W. (1978).
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
re many pala Cire Fae ; lay
(ii
The Lambert Peninsula, Ku-ring-gai Chase National Park. Physiography and the Distribution of Podzols, Shrublands and Swamps, with Details of the Swamp Vegetation and Sediments
ROBIN A. BUCHANAN
BucHANAN, R. A. The Lambert Peninsula, Ku-ring-gai Chase National Park. Physiography and the distribution of podzols, shrublands and swamps, with details of the swamp vegetation and sediments. Proc. Linn. Soc. N.S.W. 104 (1), (1979) 1980: 73-94.
The Lambert Peninsula is asymmetric, with deeply-dissected V-shaped valleys on the western side and on the eastern side extensive areas with low slope (less than 5°). Podzols, shrublands and swamps on the peninsula have been mapped using aerial photographs and ground confirmation. All podzol soils, the majority of swamps and most of the shrublands occur on areas of low slope. Areas of low slope underlain by clayey sandstone (puggy material) and extensive sandstone benches lead to poorly- drained soils and development of moist shrublands. In even less well-drained conditions swamps form, both on valley floors and valley sides. The vegetation of the swamps is divided into four types using the characteristic height and abundance of some of the larger conspicuous species. The anomalous distribution of two of the swamp species, Banksza robur and Gymnoschoenus sphaerocephalus, on the peninsula is mapped. The iron/organic-rich sediments which have accumulated in swamps to depths varying between a few centimetres and 2 metres are described. The influence of climatic fluctuations on the extent of moist shrublands and swamps is discussed.
Robin A. Buchanan, 22 Alicia Road, Mt Kuring-gaz, Australia 2080 (formerly School of Biological Sciences, Macquarie University); manuscript received 2 July 1979, accepted in revised form 21 November 1979.
INTRODUCTION
The Lambert Peninsula in Ku-ring-gai Chase National Park, on the northern edge of Sydney (Fig. 1), is an area of great beauty and botanical interest. The scenic attraction of the West Head Lookout at Broken Bay, the numerous walking trails, and the proximity to Sydney create intense pressure on the area. Yet knowledge of the biology of the area is minimal and no detailed vegetation maps or physiographic studies hitherto have been available for the area.
This paper is a first attempt to describe the general physiography of the Peninsula and to map the distribution of shrublands, of swamps, and of plant communities growing on podzolized sand accumulations on the plateau surface. An attempt is made to relate the swamps, the shrublands and the podzol communities to the physiography and geology of the peninsula. Only passing comment is made on the woodlands and forests of the area, and the vegetation on the dykes and on the Triassic Narrabeen Group is not mentioned. This group is only revealed near sea level whereas the gentle undulating plateau surface, broken by steep gullies, has been produced by weathering and erosion of the overlying Hawkesbury Sandstone.
The depositional environment of the Hawkesbury Sandstone is still under debate (Ashley and Duncan, 1977). The sedimentation units are near horizontal on a regional scale, but are commonly cross-bedded. It is an argillaceous iron-rich quartz sandstone, with the grain size commonly medium to coarse (Standard, 1969). The soils have a sandy top soil often overlying a more clayey sub-soil (Gradational Gn, or
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
74 THE LAMBERT PENINSULA
duplex Dy soils; Northcote, 1971) but in two regional surveys (Walker, 1960; Hamilton, 1976) the soils of this sandstone have been mapped as uniform sandy soils. Shale is infrequent in the Hawkesbury Sandstone sequence but on the eastern surface of the Lambert Peninsula a clayey material is quite widespread. The texture can vary from a clayey sand to clay over short (approximately 1 m) distances, both laterally and vertically, and subsurface layers of these beds are usually weathered to a pliable
ery - boundary of Salvation Creek catchment A—B sections illustrated in Fig.3
contour interval 100m
N RE a | A re (0) ] 2 km 4 BROKEN BAY West Head Lookout
(7)
g 6 CAME ERY
200 measurements
Fig. 1. Location map of the study area and a joint rose from the Salvation Creek catchment.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN 75
consistency. It is usually moist; the colour is whitish grey or, in places, mottled red to yellow. It will be referred to as puggy material.
Shale beds in the Hawkesbury Sandstone impede drainage and aid the development of swamps (Davis, 1936; Hannon, 1956; Lamy and Junor, 1965; Holland, 1974). Comparatively level areas and extensive sandstone layers can also cause deficient drainage (Pidgeon, 1938; Lamy and Junor, 1965). The term ‘swamp’ has long been applied to the wet areas around Sydney (for example by Hamilton, 1915) but more recently in Specht, Roe and Boughton (1974) the term ‘fen’ has been applied to areas dominated by Gymnoschoenus sphaerocephalus in Ku-ring-gai Chase. The traditional word ‘swamp’ is used here to describe the diverse wetlands on the peninsula.
Holland (1974) divided the swamps of the Blue Mountains (approximately 70 km west of Sydney) into two topographic types, valley-floor and valley-side, but did not name the swamps with both components. Sufficient examples of this type occur on Lambert Peninsula to warrant classifying these as a third group — composite swamps.
Pidgeon (1938) classified the swamps of the Hornsby Plateau, an area which includes Lambert Peninsula, into two vegetative types. One category was the shrub swamps, which are either classified with the driest swamp type or, more generally, grouped with moist shrubland in this paper. Pidgeon (1938) called the second type sedge swamps, and from the species list and description provided, this type covers a wide range of soil moisture. The dominant species listed typify the drier swamps on the peninsula and the large areas (up to approximately 3 ha for a single swamp) of the wetter types on the peninsula only receive a very brief mention as occurring in soaks and drainage channels. The swamps between McCarrs Creek and West Head, which were grouped as sedge swamps by Pidgeon (1938), are subdivided and described as four different swamp types.
METHODS
Aerial photographs (at a scale approx. 1:14 000) were used to locate and map shrublands, swamps and podzols and for measurement of joint orientations. The ease with which features such as vegetation boundaries and joints in rock outcrops can be seen on these photographs varies with the time since a fire. As the vegetation regrows after a fire, the early stages are insufficient to define features, and at later stages dense regrowth may obscure some detail. Aerial photographic interpretation on Lambert Peninsula is easiest approximately ten years after a fire. Identification of features was confirmed by field examination.
The base maps for Figs 1, 4 & 5 were adapted from 1:63 360 Broken Bay sheet and 1:100 000 Sydney sheet 9130. Fig. 2 was compiled from aerial photographs (Cumberland 1970) and N.S.W. 1:25 000 ortho-photomaps Broken Bay, Hornsby, Cowan and Mona Vale. Sections across the peninsula (Fig. 3) were derived from the N.S.W. Department of Lands map, Ku-ring-gai Chase 1:25 000.
All swamp profiles were measured using an Abney Level. After initial samples of subsurface layers were obtained using a two inch auger, a length of steel re-inforcing rod pushed into the ground was generally sufficient to locate sand, puggy material, swamp fill layers and the podzol pan between successive auger holes.
Botanical names are those used in Beadle, Evans and Carolin (1972) except for some recent name changes.
THE SHAPE OF THE PENINSULA
The directions of the western and eastern boundaries of the peninsula, Cowan Creek and Pittwater, and the southern boundary, Coal and Candle Creek (Fig. 1) are
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
76 THE LAMBERT PENINSULA
WW
WV _ LES & \\
a
Approx. scale
NE \ Lay VP”.
a / © OX PIS “* Z \ 5A
\ ea
wa 1g Ws
\ N
were ies LES
=<
(contour interva 10 metres)
So rock outcrop
mm cliff
ES woodland/open- forest: ** J contains podzol
Z shrubland
L__] tow woodland / woodland —-— lineament
——— track
pee swamp
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
Fig. 2. Map of the upper Salvation Creek catchment.
R. A. BUCHANAN 77
probably controlled by NE-SW and NW SE joints respectively, since these boundaries correspond to regional joint orientations (Mabbutt, 1970). Joint-control of cliffs, creeks and swamps, and vegetation boundaries is obvious throughout the peninsula and can be observed in the field and on aerial photographs. The joint rose in Fig. 1 is compiled from measurements made from aerial photographs of the Salvation Creek catchment. Joint measurements from ground survey in this catchment also show predominant NE-SW and NW-SE directions, and many features in the upper Salvation Creek area follow these orientations (Fig. 2). For example, Salvation Creek and the creek draining Swamp 11 both drain SE. The cliffs defining a trench-like valley north of Podzol 15 are orientated NW-SE and are approximately in line with a cliff and a lineament (revealed as a joint and a gap in the tree canopy along different parts of its length) in the Salvation Creek Catchment. The NE-SW orientation is less frequently exploited but one example is the cliffs near Swamp 14.
The peninsula is asymmetric about the divide running approximately N-S (Fig. 4). West of the divide it is deeply dissected by V-shaped valleys but on the eastern side, shallow basin-shaped valleys are present (Fig. 3, section A-B). For example, (Fig. 3, section C-D), the westerly flowing Yeomans Creek has deeply eroded headwaters, a middle course with a gentle slope, followed by a short steep fall to Cowan Creek. The headwaters and middle section of Salvation Creek cross a gentle easterly sloping surface, beyond which the creek falls steeply to Pittwater. Cross sections halfway along
2) _— a £ © ee 3 ‘= wn (S$) ' : 5 § 2 = % ) a a o=] P=) 3 E 7) © m a S iJ oO x o 3 3 180 (S) (7) ” 7) : | | | E 120 S >
60
<— Coal and Candle Creek
oD + Pittwater
| 0) 1 | 2 3 7 5 km 6 | | ay | 3| | © = | 5| zl =| S + | 3! =| a| ay =| m | il G | yee 120 V | 4 60 0) (0) ] km (0) 1 2km
Fig. 3. Topographic sections of the Lambert Peninsula. Sections located on Fig. 1.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
78 ; THE LAMBERT PENINSULA
these creeks (Fig. 3, sections E-F, G-H) show that the Yeomans Creek valley is steep- sided and narrow, while Salvation Creek occupies a broad valley with gently sloping sides.
Steep slopes (Slope Class 1, slope >15-20°) are virtually restricted to the coastal cliffs and some deep gullies on the eastern half of the peninsula, but on the western side these slopes intrude deeply (Fig. 4). The intermediate slope class (Class 2, 5° to 15° or 20°) covers large areas on both sides of the peninsula, but it is most common on
Slope classes :- FJ 1. > 15°- 20° N 2. 5%15° or 20° L] 3. <5°
— division between east and west drainage
SF Gymnoschoenus sphaerocephalus Gahnia sieberana
e Banksia robur
5 swamp number
5 podzol number
eal ante 0) 1 2 3km
Fig. 4. The distribution of the three slope classes, podzols, swamps and three swamp plants, Gymnoschoenus sphaerocephalus, Gahnia steberana and Banksia robur.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN 79
the western half. The most gentle slope class (Class 3, <5°) is almost confined to the eastern side, with only very small areas present in the west.
The causes of the asymmetry are not known but the gentle easterly slope of the surface of the Hawkesbury Sandstone and the easterly trend of the cross-beds (Standard, 1969) may be responsible.
PODZOLS
Podzol soils on the sandstone plateaux have only recently been described in detail (Buchanan and Humphreys, 1980) and the vegetation characteristic of these soils was first mentioned in the above paper.
All twenty-eight podzol soils (at grid references listed in Appendix 1) which were found on the peninsula occur in low slope areas (Fig. 4). Only six of these are on the western side and these six occur in isolated small areas with a gentle slope. Podzol soils develop in deep (1-4 m) quartz sand deposits which may cover an area from less than 0.1 ha to approximately 5 ha. These deposits frequently occur near creeks or swamps. The vegetation growing on these soils is very similar throughout the peninsula. The trees are usually taller (open-forest/woodland; Specht, 1970) than the surrounding Eucalyptus haemastoma Sm. association of low open-forest/low woodland. They can therefore be identified on aerial photographs with a high degree of confidence using the characteristics of gentle slope and taller vegetation, and in the field, the floristics of the tree and understorey layers aids identification of these soils (Buchanan and Humphreys, 1980).
SHRUBLANDS
The term shrubland has been selected as the clearest description of shrub- dominated areas, although it has been used in vegetation classifications (Specht, 1970; Forster, Campbell, Benson and Moore, 1977) to refer to shrub communities of specific height and density. The shrub-dominated areas on the peninsula fit into many categories in these detailed classifications and the term shrubland in this paper does not imply a specific height or density of shrubs. In a floristic survey of Ku-ring-gai Chase, Outhred, Lainson, Lamb and Outhred (in preparation) classify shrub- dominated areas in a fashion broadly consistent with that adopted by Specht.
The distribution of shrublands does not appear to be influenced by aspect or shelter (Pidgeon, 1938) but rather by soil depth and drainage. Fairly extensive areas of shrubland interspersed with rock outcrops occur on the peninsula, especially at the northern end (Fig. 5). Shrublands develop in these areas as resistant sandstone beds near the surface prevent trees becoming established. However, much of the shrubland is not broken up by rock outcrops and the presence of shrubland in these areas is controlled by a number of factors, all of which result in poorly drained soils. The majority of these moist shrublands have a low surface slope and are most common on the gentle slopes on the eastern side of the peninsula. A puggy material is often present approximately 1 m or less below the surface and it effectively prevents rapid drainage. Extensive sandstone benches also impede drainage and many moist shrublands are perched on benches with the sandstone revealed on the downslope side (Fig. 2). The extremely broken topography of the western half of the peninsula does not allow impeded drainage except on small benches.
On the eastern half, the boundary between shrubland and trees is sometimes ill- defined but it is frequently very sharp and coincides with the change from a poorly- drained to a well-drained soil. Observation during and after torrential rain showed that the surface soil under the trees never became waterlogged but the shrubland soil became waterlogged, and remained so for days. This rapid change in soil drainage
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
80 THE LAMBERT PENINSULA
may occur over as short a distance as several metres, and matches exactly the tree/shrub boundary. The most spectacular examples have a very obvious change in understorey as well, and are often the result of an abrupt junction between clayey sandstone and sandstone (Fig. 6). An equally sharp shrub and tree boundary, but a more gradual change in the understorey, takes place when the clayey sandstone thins or deepens beneath the trees.
Many of these moist shrubland/tree boundaries obviously occur at joints which mark this change in rock type and involve little or no change in surface topography.
Slope classes :- 1. > 15°- 20° 2. 5°- 15° or 20° (Gul 3° <5¢
division between east and west drainage WW shrubland interspersed with rock outcrops Z shrubland with few rock outcrops
swamp
Fig. 5. The distribution of the three slope classes; shrubland and swamps.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN 81
Joints which control the direction of cliffs often mark a change in vegetation as well as surface elevation. Fig. 2 shows a clear example of shrubland on a rock.shelf and an area covered by trees immediately below the cliff north-west of Podzol 15. Rapid changes in surface and subsurface slope can also cause an abrupt change in soil drainage.
SWAMPS
The distribution of swamps in the West Head to McCarrs Creek area is strongly controlled by lithology and topography. Areas need a fairly constant inflow of water as well as a slow loss of water for swamps to develop. The simplified and generalized longitudinal section of the swamps on the peninsula (Fig. 6) shows the suitable conditions for swamp development.
The first requirement is a gentle slope. Twenty-three of the twenty-seven mapped swamps (grid references listed in Appendix 2) occur in areas with a slope of 5° or less (Fig. 4). Only two of the valley-floor swamps, which form along creeks, occur where the general surrounding slope is greater than 5° (Nos. 17 and 21) but all of the valley- floor swamps and valley-floor elements of composite swamps have a longitudinal slope between 0° and 3°. Some valley-floor swamps are probably situated directly on extensive sandstone benches in these near horizontal segments of the creek beds. Valley-side swamps are frequently perched on rock shelves on the valley slopes and may have a slope of up to 10°.
The second requirement is the presence of the puggy material in the swamp catchment. Water percolating vertically through the overlying sandy soil and sandstone beds will saturate the puggy material and water seeping from it provides a fairly slow and constant supply to the swamps in dry periods. This layer may sometimes continue laterally under readily-draining sandstone beds and, if this is so, the catchment of these puggy layers may be considerably larger than augering indicates. The presence of this puggy material near the surface is indicated by the presence of moist shrubland, and all swamps except No. 15 and the valley-side arm of No. 4 receive drainage from moist shrubland. The swamps may occur immediately
Cnn ~ [=] sand
' t Lg tohinfer, 1 VV
4 wr UCT Va ren — ea Swamp fill
ws sh / = Clayey sandstone
ee Sandstone
Low woodland
Moist shrubland
eg. Hakea teretifolia SG RENEE ING Gan ae Swamp
e.g. Gahnia
SLVR ee it a Se LAIR ie ion . WW)
BAN ait (igi be
Fig. 6. Simplified and generalized longitudinal section of the swamps on the Lambert Peninsula.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
82 THE LAMBERT PENINSULA
downslope of such areas or they may be connected by a creek. Some swamps are directly underlain by the puggy material.
Phystographic Types Valley-floor Swamps
Valley-floor swamps are linear, as they form along creeks. They have a cross sectional slope of 0° and a longitudinal slope of 0-3°.
Swamp 21 (Fig. 7a, Fig. 9a) is an example of the simplest type of valley-floor swamp, as there is no zonation between the tall, dense swamp vegetation and the low woodland (sensu Specht, 1970) growing on the adjacent steep, rocky slopes. The iron/organic swamp fill is up to 2 m deep and a thick layer of sand is present between the swamp fill and bedrock. The main creek channel is on the least sandy side of the swamp, immediately adjacent to the rocky slope. Swamp 11 (Fig. 7b) is an example of a common valley-floor type. In the area of the section, the gradual slope change along one side of the swamp has allowed a very narrow zone of short swamp vegetation to develop between the tall and dense swamp vegetation and the low woodland. In other places along this border a thin zone of dense Banksza ertczfolia has developed and such fringes are fairly common on the peninsula. Many valley-floor and composite swamps have a short (0.5-2 m) steep sandy rise along the edge. A podzol soil supporting a distinctive podzol vegetation occurs on these rises (Buchanan and Humphreys, 1980).
Valley-side Swamps
These swamps are very variable but frequently have a fairly steep slope (3-8°) and are sometimes controlled by the underlying rock shelf. Hanging swamps described by Pidgeon (1938) are probably valley-side swamps perched on rock shelves.
Swamp 14 (Fig. 7c) is not controlled by cupping of the underlying rock shelf, as the shelf slopes upward at approximately the same angle as the soil surface, 3°-7°. No creeks drain into this swamp so water levels must be maintained by downslope seepage, and drainage is probably impeded by the relative scarcity of joints in the rock shelf. The vegetation types indicate fairly dry conditions, as only a narrow zone of dense swamp vegetation occurs. Broader zones of short vegetation occur on either side of the wettest area and the usual fringe of mixed vegetation is present along the rock shelf boundary.
Swamp 4 (Fig. 8d, Fig. 9c) is the steepest (slopes up to 14°) in the area and it is the only example of the following type. A cliff 4 m high above a puggy material layer occurs at the upslope edge of the swamp. The presence of tall and dense swamp vegetation at the base of the cliff, as well as the geological relations, indicate that water draining from the overlying sandstone seeps out of the clayey layer at the base of the cliff. Further downslope, drier conditions only enable a swamp with short vegetation to form. Sand washed down the slope has accumulated in irregularities in the bedrock, giving the hillside a smooth rather than terraced profile. The iron/organic fill forms a continuous skin over the sand and rock. On each side this hillside swamp is bordered by well-drained soils carrying a low woodland of Eucalyptus haemastoma, and the boundary between swamp and low woodland is remarkably sharp.
Composite Swamps
These swamps consist of a valley-floor element and the bordering gentle (0-3°) hillslope. A decrease in soil moisture and a greater fluctuation in soil moisture occurs from the valley-floor up to the hillslope. The water-table fluctuations are influenced by such features as the surface and bedrock slope, the permanence of the creek, the
Proc. LINN. Soc. N.S.W., 104 (1), (1979) 1980
R. A. BUCHANAN
(a) fk sandstone al SWAMP: vegetation- sb
A = puggy material ; swamp fill
—— podzol pan surface
low woodland tall and dense
—— approx. height of the vegetation ( understory in low woodland )
st
(b) 10 SWAMP:- vegetation- 8 low woodland low woodland m 6 short talland dense 4 LEO - ple as 0) 20 40 m 60 80 100 (c)
SWAMP: vegetation-
a low woodlan
8 ea
mixed
shrubland
short dense short
Tee eet we
Fig. 7. Swamp sections: a) Swamp 21, valley-floor. b) Swamp 11, valley-floor. c) Swamp 14, valley-side.
Proc. Linn. Soc. N.S.W., 104 (1), (1979) 1980
84 THE LAMBERT PENINSULA
distance from the creek, and the depth, thickness and extent of the puggy material. The type and extent of swamp vegetation on the hillslope is therefore very variable.
Swamp 13 (Fig. 8e) shows the zonation up the hillside. As usual, the tall and dense swamp vegetation is present on the valley-floor component. A change in slope from 0° to 2° and the presence of puggy material cause a rapid decrease in vegetation height to an area dominated by Gymnoschoenus sphaerocephalus. A gradual but still readily distinguishable change from short to