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PROCEEDINGS
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LINNEAN SOCIETY
NEW SOUTH WALES
VOLUME 111 (Nos 485, 486, 487 & 488; for 1989)
Sydney The Linnean Society of New South Wales 1989
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Index
Volume 111
Page Acacia iteaphylla, seedling development .... 37 Acanthodians, culmacanthid ........... 11 Achaearanea mundula ................... 25 Amino Acids, Amides, redistribution .... 37 Amphientomidae, two new species ...... 31 Anderson, G. J., seeGray, M. R......... 25 Antarctica, south eastern Australia ...... 11 ANMIECIOIONGAS 5.5 0000050000000000000008 278 Araneoidea, Theridiidae .............. 25 Argyrodes, host predation ............... 25 Argyrodes incursus sp. NOV. ............--- 25 Asteroidea, anew genus............... 293 /ASURODSCUUONGIENS © oo onc 0e 9050040040408 268 Austrochthonius australis, redescription ..... 233 Belyaevostella gen. nov. ................. 304 Belyaevostella hispida ................... 304 TBST REGIS 5 coc occossec0esesncns 67-100 Bradke, A. B., & Murray, D. R., Redistri- bution of amino acids and amides during seedling development in Acacia iteaphylla F¥. Muell. (Fabaceae: Mimo-
SOIGEAS) Mea rey sorte jens wiser ores OR eee 37 Brasimerdaes 92 bine We eae Sa eee 274 Brown, Robert, Australia 1801-5 ........ 65 Bry opliyitayen cic ssc nee ae one 82 Buchanan, R. A., Pied currawongs (Strepera
graculina): their diet and role in weed
dispersal in suburban Sydney, New
SouthsWales so oe ccs sd vee eee 241 Caenopedina alanbakert sp. nov ........... 265 Campbell, H., John Vaughan Thompson,
[By Lie Sis, eeaue tears tie ree Cee Nace nee nerece 45 Caymanostella admiranda ................ 298 Caymanostella Belyaeva, new species == GESCHIPUOM Sascha oe aes ome ctne yer: 293 Caymanostella phorcynis sp. nov. .........-. 301 Caymanostella spinimarginata ............. 298 Caymanostellidae, a review ............ 294 @lhiarophyitai pac qe5 6 ces ans cae sete eee we 77 Chernetidae, new Australian species ..... 123 Chlorophyta fie .sshqacnessncsansess 70 Chthoniidae (Pseudoscorpionida) ....... 233 Gurcipedesa acoso Are aacia wamern cin cee 53 Conicochernes doyleae sp. nOv ...........-. 123 (COMMERO NSS . 600065050000 00n00005 96 Cryptogams, Gymnosperms ........... 65 Culmacanthus antarctica sp. nov ........... 14 Culmacanthus pambulensis sp. nov ......... 17 (CHYCAGCIOD VAIS conn cccccccasccanenonge 96 IDsyormein, IPISEES .5200000c0c0a0n050000 11 Diet and role in weed dispersal in suburban
Swieleyg Air soca iyi ianyee SEN ORS Se rea 241
Echinoderm fauna checklist ............ Echinodermata, deep-water species from Norfolk Island and Wanganella Bank . Echinothunidacinn nnn panononerorncer Elix, J. A., & Streimann, H., The Lichens of Norfolk Island. 1: Introduction and the Family Parmeliaceae............
Fabaceae, Mimosoideae............... Filicopsida (True Ferns) ............... Flannery, T. F., Microhydromys musseri n. sp., a new murid (Mammalia) from the Torricelli. Mountains, Papua New Guiineare cei: oo renee nc ee JNO DARD 20262 000c000c0000000000¢ Flavoparmelia norfolkensis sp. nov. .........
Glyphodiscus mcknighti ................4. GOMASISHCES, oocccacceaccc00000008 Gray, M. R., & Anderson, G. J., A new Australian species of A7gyrodes Simon (Araneoidea: Theridiidae) which preys ONES MOSE psagis sce race aeeahatenaeaytiecee lo Groves, E. W., & Moore, D. T., A list of the cryptogams and gymnospermous plant specimens in the British Museum (Natural History) gathered by Robert Brown in Australia 1801-5 ...........
Hapalosoma pulchrum sp. nov. ........-.-- Hemuiseopsis alettae sp. nov. ............-. History, Murrary Cod Fishery .......... Holothuria (Vaneyothuria) unica sp. nov ..... Iolothuriid cae neeeeeeeeeceeeeeeneer
Kennedy, C. M. A., Redescription of Aus- trochthonius australis Hoff (Chthoniidae: Pseudoscorpionida) ...............
Kennedy, C. M. A., Conicochernes doyleae, a new Australian species of the Cherneti- dae (Pseudoscorpionida: Arachnida) .
Lake Macquarie, New South Wales ..... Lambert, M. J., & Turner, J., Redistri-
bution of nutrients in subtropical rain-
forest trees! 2. sion a Sees ae Larval silverbiddy Gerres ovatus and Gobies Weichemes: 2c sented sya eens sees ae ae Liverworts, leafy, thalloid.............. Lophostemon confertus .............-.-.-- Lycopsida “<5 ssacs adasgene an sabes acne
Maccullochella peeli .................4.. Mckenziartia, Pectocythere................ WMickenzvantiah) qucctae eee ee
257
257
263
103
37 85
215 110 110
273 273
25
233
123
PROC. LINN. SOC. N.S.W., 111(4), 1989
Mckenziartia mowbrayi sp. nov. ........... Mckenziartia portjacksonensis ............. Mckenziartia thomi sp. nov. ............-. Mesothuria (Pentchrothuria) norfolkensis sp. nov Mucrohydromys mussert sp. NOV. ........... Mikulandra, M., see Yassini,I........... Moore, D. T., see Groves, E.W. ......... Murray, D. R., see Bradke, A. B......... Murid (Mammalia), Torricelli Mountains IMNISClis.rc cea nerd yee eas HeReRSEO kor Rae ae
Nanometra duala sp. nov. ................ Neo fuscelea® inc. kssiis. wie ays 1 st Ws eSewackten wars tie Neofuscelia verrucella ................... Neothyonidium parvipedum sp. nov. ........ Nitrogen, mobilization ................ Norfolk Island, lichens ................ Norfolk Island, Wanganella Bank, north-
eastern Tasman Sea ............... Novodiniainelenach nee eee ee Nutrients, redistribution ..............
Ordovician — Silurian Stratigraphy .....
Parapanmelia A. 4.2 pent eho = ee Paraparmelia scotophylla ................. Rarmeliag yt. tse d canes: € ona boeaine MALO. GOUMIVAIS x0008000000000000000¢ Parmeliaceae, family ................. Rarmelinopsis viscevsteer ayo kes et ener: Parmelinopsis spumosa .............+4-.- IParmotremal ae Ak Sustains Hae hs SR Parmotrema austrocetratum ............... Parmotrema chinense .................-.. Parmotrema crinitum .................... Parmotrema cristiferum ..............++5. Parmotrema gardnert .................... Parmotrema rampoddense ................. Parmotrema reticulatum................... Parmotrema sancti-angelit ................ Parmotrema tinctorum .............+++.--. Pectocythere royi sp. nov. ..............-.- Pectocytheridae, Ostracoda, Crustacea .. Pedinidaers-1e) sere to eae re Pemberton, J. W., The Ordovician —
Silurian stratigraphy of the Cudgegong
— Mudgee District, New South Wales Rentacnimusscuropacus ene ee Eee Percichthyidaeneae eee ee Phacophytacastiei Moe oe ee eee ee See Pied Currawongs (Strepera graculina) ...... Phyllophonicdaceen eee eenneraereeee iRiscess Devonian) eee eno ROLY 20a: oro ther it ee eee Pressey, R. L., Wetlands of the lower
Clarence Floodplain, northern coastal
INERT SOWEN WANES 50c00cc0cg0cd00c0 Pressey, R. L., Wetlands of the lower
Macleay Floodplain, northern coastal
New South Wales .................
PROC. LINN. SOC. N.S.W., 111(4), 1989
143
157
INDEX
Pseudoscorpionida, Arachnida .........
Psilopsidad i. < snc cers one sce ee one Psocoptera, Insecta ................... Pteridophyta: <0.) son oanusee Gea es Rainforest trees, subtropical ........... Rhodophyta’ <:¢4.5..24.52.25 See ee Rowe, F. W. E., A review of the family
Caymanostellidae (Echinodermata:
Asteroidea) with the description of a new species of Caymanostella Belyaev and a new genus ................... Rowe, F. W. E., Nine new, deep-water species of Echinodermata from Norfolk Island and Wangangella Bank, north- eastern Tasman Sea, with a checklist of the echinoderm fauna.............. Rowland, S. J., Aspects of the history and fishery of the Murray Cod Maccul- lochella peels (Mitchell) (Percichthyidae)
Seopsis incisa sp. NOV. ........-++-+--+-0 SiltstonesyAztee Sam aa sane eee cee Smithers, C. N., Two new species of
Amphientomidae (Insecta: Psocop- tera), the first record of the family for Australia: fcc ccsteccius ten sree
Steffe, A. S., Tidal and diel variations in the abundance of larval fishes in Botany Bay, New South Wales, with emphasis on larval silverbiddy Gerres ovatus (Fam. Gerreidae) and gobies (Fam. Gobiidae) ee. Cae ee eae
Stratigraphy, Cudgegong-Mudgee District
Streimann, H., see Elix, J. A. ...........
Synallactidae: . 2.45. JE. a
Tethyaster tangaroae sp. nov. .............. Theridiidae, Araneoidea .............. Turner, J., see Lambert, M.J............
Variations in abundance, tidal, diel, Botany Bay) oo. cretistameatsteeoe
Weed dispersal, currawongs ............ Wetlands lower Clarence floodplain ..... Wetlands lower Macleay floodplain ......
Xanthopanneliaaeeen eee er ee Xanthoparmelia amplexula ............... Xanthoparmelia australasica ..............
Yassini, I., & Mikulandra, M., Mckenziartia and Fectocythere (Pectocytheridae, Ostra- coda, Crustacea) in Lake Macquarie, New South Wales .................
Young, G. C., New occurrences of culmacanthid acanthodians (Pisces, Devonian) from Antarctica and south- eastern Australia ..................
293
31
131
PROCEEDINGS of the
LINNEAN SOCIETY
NEW SOUTH WALES
VOLUME 111 NUMBERS 1-4
NATURAL HISTORY IN ALL ITS BRANCHES
THE LINNEAN SOCIETY OF NEW SOUTH WALES
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Honorary Editors: T. G. VALLANCE — Department of Geology & Geophysics, ‘University of Sydney, Australia, 2006. (Numbers 1 & 2.) J. R. MERRICK — Graduate School of the Environment, Macquarie University, North Ryde, N.S.W., Australia, 2109. (Numbers 3 & 4.)
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Cover motif: The gastropod collected ‘16 miles cast of Wollongong’ and described by Charles Hedley as Stiva ferruginea gen. ct sp. nov. (Proc. Linn. Soc. N.S.W. 29, 1904: pl. LX, no. 23).
PROCEEDINGS of the
LINNEAN SOCIETY
NEW SOUTH WALES
VOLUME 111 NUMBER 1
Redistribution of Nutrients in Subtropical Rainforest ‘Irees
MARCIA J. LAMBERT and JOHN TURNER
(Communicated by D. W. EDWARDS)
LAMBERT, M. J., & TURNER, J. Redistribution of nutrients in subtropical rainforest trees. Proc. Linn. Soc. N.S.W. 111 (1), 1989: 1-10.
Fresh foliage and leaf litterfall from trees and understorey plants in a N.S.W. sub- tropical rainforest were chemically analysed to estimate nutrient redistribution. In general, the proportion of nutrients redistributed in these species at time of leaf litterfall is low. Results available on nutrient redistribution from the same species in this area during heartwood formation showed that trees which redistributed phosphorus from foliage, redistributed little from heartwood and vice-versa. By way of contrast, species in sclerophyll forests were highly efficient at nutrient redistribution from both leaves and wood. Oo me peters
Marcia J. Lambert and John Turner, Forestry Commission of N.S.W., PO) Bil aomh BeeBibl datcal | air tralia 2119; manuscript receved 19 November 1986, accepted for publication 23 TES eG f.
| IBRARY INTRODUCTION j 1 Nutrient cycling within forests is critical for long term rdaintenatol bos produlgo ty
and stability. It involves nutrient uptake, utilization and accumulation by vegetation, together with the return of nutrients to the soil through litterfall, leaching and root sloughing. Comparisons of nutrient cycles involve considerations HK Spe iesrelable ss. to obtain nutrients from soils with low nutrient status and then rétain-the-nutrients.___ within systems (Turner, 1975). In order to assess turnover of nutrients, various indices have been developed. For example, turnover of litter on forest floor has been compared
by using a ‘k’ factor which relates the input of litter (L) to the mass of litter (F) on the forest floor. The ‘k’ factor = L/F and assumes a steady state forest floor weight (Jenny e¢
al., 1949; Olson, 1963; Richards and Charley, 1977) and gives an indication of the rate of
loss (decomposition) by the litterfall in relation to accumulation on the forest floor.
Relative efficiency of nutrient acquisition from soil, nutrient utilization requirements and efficiency of redistribution of nutrients can be assessed within the tree component of an ecosystem. These comparisons are difficult, but some can be made. For example, an index of nutrient utilization within a tree is often obtained using foliage nutrient con- centrations (Lambert and Turner, 1983; Lambert et a/., 1983). Nutrient redistribution may be estimated as withdrawal of nutrients, both during leaf abscission (Ashton, 1976; Attiwill, 1980; Turner and Lambert, 1983) and in heartwood formation (Lambert, 1981). Such estimates have been made in only a few forests in Australia and have been carried out predominantly in stands dominated by a single species (Hingston et al., 1979; Attiwill, 1980; Turner and Lambert, 1983) where monthly leaf litterfall data were com- pared with those for live leaf material on the trees. In studies of forest stands including a variety of species, and particularly in conditions where organic matter decomposition and tissue leaching can be quite rapid, the use of monthly litterfall samples becomes inappropriate.
Subtropical rainforests are associated with relatively fertile soils (Baur, 1957; Webb, 1969; Lambert et a/., 1983) whereas eucalypt forests are on soils with much lower fertility (Baur, 1957; Webb, 1969; Turner and Kelly, 1981). Subtropical rainforests (Baur, 1965) have high species diversity with often in excess of 30 species ha" in the overstorey. They are notable in northern N.S.W. for the absence of Eucalyptus species, a genus which
PROG. LINN. SOC. N.S.W., 111 (1), 1989
2 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES
dominates most other coastal and tableland forest types in N.S.W. During a programme of study in a subtropical rainforest located on the New South Wales Border Ranges, leaf material was sampled from a range of species 1n order to obtain indices of nutrient distri- bution and cycling patterns within this forest. Fresh litter was specifically sampled to provide estimates of nutrient redistribution in various species. These data were com- bined with differences in heartwood and sapwood nutrient concentrations and compared with similar data from coastal sclerophyll forests.
STUDY SITE
The study site was originally described by Lambert e¢ al. (1983) and was located in forests of the Border Ranges (153°E, 28°38’S), west of Murwillumbah. The altitudinal range of the forest is 600-1200m above sea level. Annual rainfall is 3000mm. Plots were selected from within subtropical rainforest growth experiments (Burgess et al., 1975; Horne and Gwalter, 1982). Soils from the region are derived from Tertiary volcanic rocks of the Mt Warning Shield and are predominantly of basaltic composition (Stevens, 1976). The basalts have given rise to kraznozems, that is, deep well-structured red clay loams with clay sub-soils having a relatively uniform appearance and depth (Beckman and Thompson, 1976). The soils are high in nutrients, particularly when compared with soils supporting sclerophyllous species (Lambert et al., 1983).
METHODS
Trees were sampled for foliage and wood during logging operations near the study site. Within a species, nutrient variability in foliage concentration was found to be low (Lambert et al/., 1983; Lambert and Turner, 1986). Foliage samples were bulked from within the crown but only fully-formed leaves were used; that is, very young or damaged leaves were omitted. Samples were placed in paper bags, oven dried at 70°C, ground and analysed for various chemical elements (Lambert, 1983). The results for overstorey and understorey trees have been reported elsewhere. Wood discs were taken from the stem approximately 1m above the ground. These were air dried and then separated into bark, sapwood and, where present, heartwood. The individual components were ground and analysed for the same chemical elements as the foliage samples (Lambert, 1983).
Freshly-fallen leaf samples were sampled beneath the crown of selected species within the research plot and were those recently fallen so that there was minimal effect due to decomposition and leaching. Where leaves were caught on understorey vegeta- tion, they were preferentially selected so that ground contact was minimized. Acquiring sufficient suitable material of many species was difficult and hence single, bulked samples were used rather than replicates. The leaf litterfall samples were dried, ground and analysed in the same way as foliage samples.
A list of the common names, scientific names and authorities of species sampled on the study site is given in the Appendix.
RESULTS
Results of the analyses are presented in detail in Table 1; trends are summarized in Table 2. Changes in concentration of nitrogen and phosphorus between overstorey leaf and litterfall were relatively minor in most cases. There were obvious exceptions, such as Dendrocnide excelsa, Solanum aviculare and Solanum mauritianum, these tending to be very high in nutrients initially. The other exception was Orites excelsa which has relatively low requirements for both phosphorus and nitrogen and appears to be also efficient at retranslocation, that is, there is high nutrient removal by retranslocation even in a situ-
PROC. LINN. SOC. N.SW., 111 (1), 1989
M. J. LAMBERT AND J. TURNER
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NUTRIENT REDISTRIBUTION IN RAINFOREST TREES
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PROC. LINN. SOC. N.S.W., 111 (1), 1989
M. J. LAMBERT AND J. TURNER 9)
ation of high nutrient availability (Kelly e¢ a/., 1983). Calcium typically increased in con- centration in the litter of most species, an exception being Orites excelsa. The patterns of other elements were much more irregular (Table 2). Potassium and manganese tended to decline and accumulate respectively. Caldcluvia paniculosa and Orites excelsa accumu- lated aluminium in the foliage but the concentration of this element was lower in the litter, the Orites excelsa to a higher degree possibly showing different patterns of handling an antagonistic element. Of the 48 samples assessed, heartwood was detectable in only 12 and in these instances relatively few had significant retranslocation of nutrients (Lambert et al., 1983).
Within the non-woody understorey species, nutrients were retranslocated between young and old tissues as in the woody species (Table 3). Cyathea australis, for example, redistributed 45% of phosphorus in the aging of fronds. Archontophoenix cunninghamiana was an efficient retranslocater of phosphorus (65%) and potassium (96%), although there may be a higher proportion leached as it is difficult to estimate when foliage of this species can be classified as litterfall as the older foliage hangs down next to the stem. Asplenium nidus retranslocated much smaller quantities during senescence.
DISCUSSION
Estimates of nutrient retranslocation are relative measures and to understand the ecological significance of nutritional patterns, comparisons with other forest types have been used. Within the subtropical rainforest, retranslocation of nutrients within foliage prior to abscission appears to be relatively low for most overstorey species, however, exceptions were: Dendrocnide excelsa, this species having fairly high nutrient require- ments; Orites excelsa, which is an aluminium-accumulating species and appears to have an efficient system of retranslocation; and two solanaceous species (Solanum aviculare and Solanum mauritianum which invade and grow immediately after soil disturbance and live for only a relatively short time. It appears they have both high nutrient demands and efficient retranslocation. The level of retranslocation may be compared with that in sclerophyllous species and coachwood which generally grow on nutritionally poorer soils than the subtropical rainforest (Table 4). In these species, phosphorus, nitrogen, mag- nesium and potassium are all retranslocated, while calcium and aluminium are accumulated. The calcium pattern appears typical of mature foliage in many trees. Further, there were consistently high removals of nutrients in these species during heart- wood development. The pattern of heartwood development and retranslocation was either absent or very low in the majority of species sampled in this rainforest.
Generally, nutrient redistribution is an important component, along with uptake, in fulfilling forest stand nutrient requirements (Turner and Lambert, 1983). However, if the general principle is that the subtropical rainforest tree species generally redistribute nutrients at a low level, whereas the sclerophyllous and cool temperate rainforest species are more efficient at redistribution, is this a function of generally higher soil nutrient availability? Further, iflarge fertilizer quantities were applied to eucalypt species or they were located on more fertile soils, would heartwood production be reduced or litter retranslocation lowered? Eucalypts growing naturally across a range of fertilities do not have significantly different foliage nutrient concentrations (Lambert and Turner, 1983) or nutrient redistribution patterns. This suggests that soil nutrients are not controlling the patterns but rather that they are evolutionarily determined. The reverse situation of growing subtropical rainforest trees on poor sites is harder to test, mainly because it appears that soil nutrients are delineators of species (Turner and Kelly, 1981) and hence rainforest species are out-competed on poor sites.
Within the subtropical rainforest there were some species such as Orites excelsa
PROG. LINN. SOG. N.S.W., 111 (1), 1989
NUTRIENT REDISTRIBUTION IN RAINFOREST TREES
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PROC. LINN. SOC. N.S.W., 111 (1), 1989
8 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES
TABLE 4
Published information for various Australian forest species on redistribution from foliage during abscission
Forest Reference
Type
IP Ca Meg K Al (ppm)
Wet Sclerophyll E. grandis — foliage 895 4800 2690 6450 80 Turner and — litter 340 7065 2160 1955 230 Lambert — change -0.61 =599 +2265 -530 -4495 +150 | (1983) E. regnans* — fohage 0.79) 780 5730 5730 Ashton — litter 0.87 300 5450 1240 (1976) — change (0), 112 -480 -280 -4490) E. pilularis — foliage 1.21 620 3420 3465 3810 120 | Turner and — litter 070 300 WAI ASSO. lleXO) 330 | Kelly — change -0.51 -320 +9190 -1075 -2670 +210 | (1981) Lophostemon — foliage 1.61 1205 V525) 2735 11035 195 Turner and confertus — litter 0.81 529) 12220 2580 1750 820 | Kelly — change -0).80 -680 +2695 159 -9285 +625 | (1981) Rainforest Ceratopetalum — foliage 1.10 505 10365 2990 4540 TOUS Turner and apetalum = litter 0.71 210 7890 1740 1030 8200 | Kelly — change =), 39 -295 -27495 -1250 -3510 +925 (1981) Nothofagus — foliage 1010 Ashton cunningham — litter _ 300 (1976) — change -710 Athosporia — foliage 1500 Ashton moschatum — litter 890 (1976) — change -630 Dry Sclerophyll E. sveberi — foliage 440 Ashton — litter 110 (1976) — change -330
4 Qnd year leaf and leaf litter analyses for a mature stand.
which gave a similar pattern to the sclerophyllous pattern of redistribution. Whether this is an indication that the species had evolved on poorer soils and now survive within the subtropical rainforest is difficult to ascertain, but it is suggested that the com- bination of high litter and heartwood redistribution efficiencies, particularly for phos- phorus, is indicative of evolution on soils with low nutrient status.
A further question arises as to the relationship between redistribution during leaf abscission and that during heartwood formation for the rainforest species. For most nutrients there was no relationship, but for phosphorus and nitrogen, there was an inverse relationship (Fig. 1). That is, as the proportion of phosphorus redistributed in leaves increased, the proportion redistributed in heartwood formation or its formation at all, decreased. The exception to this was Orites excelsa which, as noted above, tends to have unusual patterns of nutrient utilization compared with other species.
The general pattern for phosphorus in rainforest trees, is that as phosphorus becomes more efficient at redistribution in one type of tissue, it becomes less efficient in another. The species which most noticeably form heartwood are not redistributing during leaf senescence. This is the opposite pattern to that found in Eucalyptus and other sclerophyllous species (Fig. 1) where there is simultaneous redistribution from leaf litter
PROG. LINN. SOC. N.S.W., 111 (1), 1989
M. J. LAMBERT AND J. TURNER
Ko)
ROSE RUS" fe aa 80 60 40 20
-40 -20 0 20 40 60 LEAF LITTER REDISTRIBUTION (%)
Fig. 1. Relationship between redistribution of phosphorus during heartwood formation and leaf abscission (¢ — rainforest species; x — eucalyptus and sclerophyllous species; L] — Ceratopetalum apetalum; O — Orites excelsa).
and heartwood. Orites excelsa relates closely to the ‘sclerophyllous’ pattern which includes E. pilularis, E. dives, E. maculosa, E. rossu, E. rubida, E. obliqua, E. grandis, Lophostemon confertus and Casuarina torulosa, these being the species where mature green leaf, leaf litterfall, sapwood and heartwood concentrations were available. The phosphorus re- distribution pattern in coachwood, however, was similar to that found generally in the subtropical rainforest species even though this species grows in differently structured rainforest (Baur, 1965).
Comparisons with other species (Table 4) were based on results for mature leaves on the tree and fresh litter using comparable sampling techniques. Fully-developed younger leaves have nutrient concentrations different to those in older leaves, so that there are different patterns of redistribution taking place within the crown. Further, there are different patterns between different-aged forests of the same species (Ashton, 1975). The pattern for nitrogen was similar to that for phosphorus for the rainforest species. Orites excelsa was again found to form a different pattern. Nitrogen was generally not as efficiently redistributed as phosphorus during heartwood formation. Ceratopetalum apetalum and Lophostemon confertus in this case followed the rainforest pattern, while the eucalypts were differently distributed.
While for most nutrients there are relationships between foliage litter and heart- wood redistribution which may be described as specific to species/site, phosphorus and nitrogen have given separate patterns of nutrient cycling. In rainforests, where phos- phorus and nitrogen are usually readily available, a certain amount of ‘energy’ is apparently expended in retaining nutrients in biomass and hence not all tissues are affected. In the case of the lower phosphorus sclerophyllous forest, all available phos- phorus is apparently redistributed, this being the primary limiting nutrient. The pattern of Orites excelsa possibly indicated that it evolved in a low phosphorus environment.
References ASHTON, D. H., 1976. — Phosphorus in forest ecosystems at Beenak, Victoria. /. Ecol. 64: 171-186. ATTIWILL, P. M., 1980. — Nutrient cycling in a Eucalyptus obliqua (L-Hérit.) forest — IV. Nutrient uptake and nutrient return. Aust. J. Bot. 28: 199-222.
PROG. LINN. SOC. N.S.W., 111 (1),
10 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES
Baur, G. W., 1957. — Nature and distribution of rain-forests in New South Wales. Aust. J. Bot. 51: 190-222.
— , 1965. — Forest types in New South Wales. For. Comm. N.S.W. Res. Note No. 17.
BECKMAN, G. G., and THOMPSON, C. H., 1976. — The soils. In The Border Ranges — a land use conflict in regional perspective (cds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Qucensland.
BurGess, I. P., FLoyp, A., KikKAwA, J., and PATTIMORE, V., 1975. — Recent developments in the silvi- culture and management of subtropical rainforest in N.S.W. Proc. Ecol. Soc. Aust. 9: 74-84.
HINGSTON, F. J., TURTON, A. G., and DIMMOCK, G. M., 1979. — Nutrient distribution in Karri (Eucalyptus diversicolor F. Muell.) ecosystems in southwest Western Australia. For. Ecol. Managem. 2: 133-158. HORNE, R., and GWALTER, J., 1982. — The recovery of rainforest overstorcy following logging. I. Subtropi-
cal rainforest. Aust. For. Res. 13: 29-44. JENNY, H., GesseL, S. P., and BINGHAM, F. T., 1949. — Comparative study of decomposition rates of organic matter in temperate and tropical regions. Soz/ Scz. 68: 419-432. KELLY, J., LAMBERT, M. J., and TURNER, J., 1983. — Available phosphorus forms in forest soils and thcir possible ecological significance. Commun. Soil Sci. Plant Anal. 14: 1217-1234. LAMBERT, M. J., 1981. — Inorganic constituents in wood and bark of New South Wales forest tree species. For. Comm. N.S.W., Res. Note 45. 43 pp. —., 1983. — Methods for chemical analysis. For. Comm. N.S.W. Tech. Pap. 25. Third Edition. 187 pp. , and TURNER, J., 1983. — Soil nutrient-vegetation relationships in the Eden area, N.S.W. III. Foliage nutrient relationships with particular reference to Eucalyptus sub genera. Aust. For. 46: 200-209. , and , 1986. — Nutrient concentrations in foliage of species within a New South Wales sub-tropical rainforest. Ann. Bot. 58: 465-478. . , and KELLY, J., 1983. — Nutrient relationships of tree species in a New South Wales sub-tropical rainforest. Aust. For. Res. 13: 91-102. OLSON, J. S., 1963. — Energy storage and the balance of producers and decomposers in ecological systems. Ecology 44: 322-331.
?
RICHARDS, B. N., and CHARLEY, J., 1977. — Carbon and nitrogen flux through native forest floors. In Nutrient cycling in indigenous forest ecosystems: 65-81. Perth: C.S.I.R.O. Division of Land Research and Management.
STEVENS, N. C., 1976. — Geology and landforms. In The Border Ranges — a land use conflict in regional perspective (eds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Queensland. TURNER, J., 1975. — Nutrient cycling in a Douglas-fir ecosystem with respect to age and nutrient status. Seattle, Washington: University of Washington, Ph.D. thesis, unpubl. , and KELLY, J., 1981. Relationships between soil nutrients and vegetation in a north coast forest, New South Wales. Aust. For. Res. 11: 201-208. , and LAMBERT, M. J., 1983. — Nutrient cycling within a 27-year-old Eucalyptus grandis plantation in New South Wales. For. Ecol. Managem. 6: 155-168. WEBB, L. J., 1969. — Edaphic differentiation of some forest types in eastern Australia. II. Soil chemical factors. J. Ecol. 57: 817-830.
APPENDIX
List of common names, scientific names and authorities of species from the study site
Common name
Bangalow palm Birds nest fern
Botanical name
Archontophoenix cunninghamiana (H. Wendl.) H. Wend. et Drude Asplenium nidus L.
Black booyong Heritiera actinophylla (F. M. Bail.) Kosterm. Bollygum Neolttsea reticulata (Meisn.) F. Muell. Brushbox Lophostemon confertus (R. Br.) Peter G. Wilson e J. T. Waterhouse Coachwood Ceratopetalum apetalum D. Don
Corkwood Caldcluvia paniculosa (F. Muell.) Hoogl. Doughwood Euodtia micrococca F. Muell.
Giant stinging tree Dendrocnide excelsa (Wedd.) Chew Kangaroo apple Solanum aviculare Forst. f.
Lace fern Nephrolepis sp.
Pigeonberry ash Cryptocarya erythroxylon Maiden et Betche Prickly ash Orites excelsa R. Br.
Red carabeen Geissois benthamiana F. Muell.
Rosewood Dysoxylum fraseranum (A. Juss.) Benth. Teak Flindersia australis R. Br.
Tree fern Cyathea australis (R. Br.) Domin
White booyong Heritiera trifoliolata (F. Muell.) Kosterm.
Wild tobacco tree Yellow carabeen
Solanum mauritianum Scop. Sloanea woollsii F. Muell.
PROC. LINN. SOC. N.S.W., 111 (1), 1989
New Occurrences of Culmacanthid Acanthodians (Pisces, Devonian) from Antarctica and southeastern Australia
G. C. YOUNG
YOUNG, G. C. New occurrences of culmacanthid acanthodians (Pisces, Devonian) from Antarctica and southeastern Australia Proc. Linn. Soc. N.S.W. 111 (1), 1989: 11-24.
Two new spccies of the acanthodian fish Culmacanthus Long 1983 are described from the lower part of the Aztec Siltstone of southern Victoria Land. Antarctica, and Facies 3 of the Boyd Volcanic Complex of southeastern New South Wales. Both are represented only by cheek plates. That of C. antarctica sp. nov. has distinctive ornament, a longer lateral than dorsal lamina, and the infraorbital sensory groove passing off the ventral margin of the plate. C. pambulensis sp. nov. resembles the type species C. stewarti Long in the shape of the posterior margin of the check plate and the ventral course of the infraorbital sensory canal, but differs in its proportions, the shape of the anterior margin, and the much finer dermal ornament. C. antarctica sp. nov. is considered to be the oldest (?late Middle Devonian) because it is associated with turiniid thelodont scales. The two other species of Culmacanthus occur with bothriolepid and phyllolepid placoderms in assemblages considered to be early Late Devonian (Frasnian). The specific differences described herein suggest however that they are not precise cor- relatives. Culmacanthid acanthodians are only known from southeastern Australia and southern Victoria Land, a distribution pattern previously noted in chondrichthyans and placoderms from the same faunas, and suggesting that Culmacanthus was an cast Gondwanan endemic.
G. C. Young, Division of Continental Geology, Bureau of Mineral Resources, PO. Box 378, Canberra, Australia 2601; manuscript received 15 December 1987, accepted for publication 24 August 1988.
INTRODUCTION
The acanthodians are a major group of Palaeozoic gnathostome fishes, with a fossil record from Silurian to Permian. During the Devonian Period they were widely dis- tributed in both non-marine and marine aquatic environments, and their tiny scales and characteristic fin-spines are commonly represented in microvertebrate assem- blages. However articulated specimens are much less common, and knowledge of acanthodian morphology is based mainly on a few localities in Europe and North America which have yielded well-preserved complete specimens (for a comprehensive review of the group see Denison, 1979).
A. S. Woodward was the first to describe Australian acanthodians, from the Lower Carboniferous Mansfield group of Victoria (Woodward, 1906), and he also published the first descriptions of the group from Antarctica (Woodward, 1921). Isolated acantho- dian scales and spines occur widely in Devonian rocks of eastern and central Australia (Fig. 1), and in recent years some well-preserved articulated acanthodians have been described from southeastern Australia (e.g. Long, 1983a, 1986a). The best-known locality is Mount Howitt in east central Victoria, which is the type locality for the genus Culmacanthus Long 1983, of which additional material is described in this paper. With these two new species the genus Culmacanthus is now known from three localities in southeastern Australia, and one locality in Antarctica (Fig. 1). Its distribution pattern matches that seen in several other taxa of Devonian fishes, the palaeogeographic sig- nificance of which is discussed below. All described specimens are housed in the Com- monwealth Palaeontological Collection (prefix CPC), Bureau of Mineral Resources,
PROC. LINN. SOC. N.S.W., 111 (1), 1989
12 DEVONIAN FOSSIL FISHES
abs Als MR WC 4 A ARS AUSTRALIA
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20/09/172
Fig. 1. Reconstruction of East Antarctica against Australia, modified from the Gondwana reconstruction of Lawyer and Scotese (1987). showing the main localities of Devonian acanthodian fishes in the southwest Pacific region. Numbered localities are the only known occurrences of culmacanthid acanthodians, as dealt with in this paper; 1, Mount Crean, Lashly Range, southern Victoria Land (Culmacanthus antarctica sp. nov.);