aa ag ta aS ae IEA oo coe fee os Pers tre abe es ar ee erictys at 48 oh ah ae pes hst Ad payee 4 Be eieatanes =e 13 a ae a i as: et PROCEEDINGS OF THE LINNEAN SOCIETY NEW SOUTH WALES VOLUME 111 (Nos 485, 486, 487 & 488; for 1989) Sydney The Linnean Society of New South Wales 1989 sayviasaDodt * « "aM 46) wi zm a i S be a. ST IAW HTUOe Wa © -_ y as Ay eh: u te Oe ee | 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 Founded 1874. Incorporated 1884. The Society exists to promote ‘the Cultivation and Study of the Science of Natural History in all its Branches’. It holds meetings and field excursions, offers annually a Linnean Macleay Fellowship for research, contributes to the stipend of the Linnean Macleay Lecturer in Micro- biology at the University of Sydney, and publishes the Proceedings. Meetings include that for the Sir William Macleay Memorial Lecture, delivered biennially by a person eminent in some branch of Natural Science. Membership enquiries should be addressed in the first instance to the Secretary. Candidates for election to the Society must be recommended by two members. The present annual subscription is $35.00. The current rate of subscription to the Proceedings for non-members is set at $50.00 per volume. Back issues of all but a few volumes and parts of the Proceedings are available for purchase. A price list will be supplied on application to the Secretary. OFFICERS AND COUNCIL 1988-89 President: T. G. VALLANCE Vice-presidents: P. M. MARTIN, HELENE A. MARTIN, C. N. SMITHERS, Honorary Treasurer: 1. G. PERCIVAL Secretary) BARBARA J. STODDARD Council: A. E. J. ANDREWS, T. C. CHAMBERS, JUDITH H. K. EASTMAN, M. R. GRAY, SUSAN J. HAND, D. S. HORNING, L. A. S. JOHNSON, R. J. KING, HELENE A: MARTIN, P. M. MARTIN, J. Ri: MERRICK, 2a IMINO SCOUT sl SG MEINE | IPO IDL, vhs UNINC S08. C. N. SMITHERS, T: G. VALLANCE, KAREN L. WILSON Honorary Editors: T. G. VALLANCE — Department of Geology & Geophysics, ‘University of Sydney, Australia, 2006. (Numbers 1 & 2.) J. R. MERRICK — Graduate School of the Environment, Macquarie University, North Ryde, N.S.W., Australia, 2109. (Numbers 3 & 4.) Linnean Macleay Fellow: JULIA M. HUSH Linnean Macleay Lecturer in Microbiology: K.Y. CHO Auditors: W. SINCLAIR & CO. The postal address of the Society is PO. Box 457, Milson’s Point 2061, N.S.W., Australia. Telephone (02) 929 0253. © Linnean Society of New South Wales 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 G6t- gI+ L8+ 6+ CO@t GGT Ogzt O8bS- ObI+ OL- 2E 0- osueyo — CHIT 09 OFI 0G OISE O€I% O6II O8EF C8 O64 28°9 Jy — OF9I PF 6G Ib GOSE S822 096 0982 CILb OIOST tL jeap — DUDIULDY]UAG S210SS194) OSg+ 2 Og+ GI- OSE+ G89F- OSbI+ O29Z- O09T+ ZGII- 690+ osueyo — 08 (4 OI Ol OSS GOIZ 06SZ O€9I OZLI €L0 8's Joy — 0¢ 2 OL GZ ae 06Z9 OFIT 00EF 09SI 6229 €G'b jesp — Ds]29% Sa}UUC) OOSI- Zit ZOl+ OT 06>- O8Zt+ sGEt GZ8S- GzG- cgogt Z1'0- osueyo — 098 CE COI ¢9 066 OOF] Scr 0969- -S88I O6IZI IgG Jou — 0962 6% 69 CL 080I 029 aa C8IZI OIZ CCI6 60'S Jeop — uajpdxosyjdsa vdsns0jgy Gye Ber Gir C- 0Z+ 00Z- —-09- OST9- 09I- OO0FT+ 661+ osueyo — GEEG 0G OFI GZ CEZ a! 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LINN. SOC. N.S.W., 111 (1), 1989 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES Gpet G+ Geer Gis OG Gina OIE CoG= GI GGe OR |) NO» Gi aI) — GeGn a At OST GI 018 G88. «G8 OPIS GOLb = O06PT = GL 260 99°9 tony = Q6Sh Gt, G9 0¢ 096 OE! COOL 6208 (66% S626 GGL OVI 0€ + jeay = 21s7j00M vauvo]y Gobi —GIE Cone GCQO- OF = GAS + GrLo- 00h O0S0G- OFT 600° — 88'0- osurys — OI+I GG OGI 02 GIG COT OGG OLY6 G6E% SF69 0L6 G6 26h puy = OGLE — LC GG GZ OLIN GG GLE GIZZI G6L6 S668 OWT 861 02'S eop DIDpONOfIAT Ds ITAA FY OLb- GI Q9e+ OZ+ OGt OFZ+ “G6I OLA fe i 0 a = is = A se Na, ee ee Ie, ELLE AO a tt = + ee = BL + 0 a 0 OD OAO OOO NO O!0 TOnGro, 096 00 O46. 0 Gu Ble. 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N.S.W., 111 (1), 1989 8 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES TABLE 4 Published information for various Australian forest species on redistribution from foliage during abscission Forest Reference Type IP Ca Meg K Al (ppm) Wet Sclerophyll E. grandis — foliage 895 4800 2690 6450 80 Turner and — litter 340 7065 2160 1955 230 Lambert — change -0.61 =599 +2265 -530 -4495 +150 | (1983) E. regnans* — fohage 0.79) 780 5730 5730 Ashton — litter 0.87 300 5450 1240 (1976) — change (0), 112 -480 -280 -4490) E. pilularis — foliage 1.21 620 3420 3465 3810 120 | Turner and — litter 070 300 WAI ASSO. lleXO) 330 | Kelly — change -0.51 -320 +9190 -1075 -2670 +210 | (1981) Lophostemon — foliage 1.61 1205 V525) 2735 11035 195 Turner and confertus — litter 0.81 529) 12220 2580 1750 820 | Kelly — change -0).80 -680 +2695 159 -9285 +625 | (1981) Rainforest Ceratopetalum — foliage 1.10 505 10365 2990 4540 TOUS Turner and apetalum = litter 0.71 210 7890 1740 1030 8200 | Kelly — change =), 39 -295 -27495 -1250 -3510 +925 (1981) Nothofagus — foliage 1010 Ashton cunningham — litter _ 300 (1976) — change -710 Athosporia — foliage 1500 Ashton moschatum — litter 890 (1976) — change -630 Dry Sclerophyll E. sveberi — foliage 440 Ashton — litter 110 (1976) — change -330 4 Qnd year leaf and leaf litter analyses for a mature stand. which gave a similar pattern to the sclerophyllous pattern of redistribution. Whether this is an indication that the species had evolved on poorer soils and now survive within the subtropical rainforest is difficult to ascertain, but it is suggested that the com- bination of high litter and heartwood redistribution efficiencies, particularly for phos- phorus, is indicative of evolution on soils with low nutrient status. A further question arises as to the relationship between redistribution during leaf abscission and that during heartwood formation for the rainforest species. For most nutrients there was no relationship, but for phosphorus and nitrogen, there was an inverse relationship (Fig. 1). That is, as the proportion of phosphorus redistributed in leaves increased, the proportion redistributed in heartwood formation or its formation at all, decreased. The exception to this was Orites excelsa which, as noted above, tends to have unusual patterns of nutrient utilization compared with other species. The general pattern for phosphorus in rainforest trees, is that as phosphorus becomes more efficient at redistribution in one type of tissue, it becomes less efficient in another. The species which most noticeably form heartwood are not redistributing during leaf senescence. This is the opposite pattern to that found in Eucalyptus and other sclerophyllous species (Fig. 1) where there is simultaneous redistribution from leaf litter PROG. LINN. SOC. N.S.W., 111 (1), 1989 M. J. LAMBERT AND J. TURNER Ko) ROSE RUS" fe aa 80 60 40 20 -40 -20 0 20 40 60 LEAF LITTER REDISTRIBUTION (%) Fig. 1. Relationship between redistribution of phosphorus during heartwood formation and leaf abscission (¢ — rainforest species; x — eucalyptus and sclerophyllous species; L] — Ceratopetalum apetalum; O — Orites excelsa). and heartwood. Orites excelsa relates closely to the ‘sclerophyllous’ pattern which includes E. pilularis, E. dives, E. maculosa, E. rossu, E. rubida, E. obliqua, E. grandis, Lophostemon confertus and Casuarina torulosa, these being the species where mature green leaf, leaf litterfall, sapwood and heartwood concentrations were available. The phosphorus re- distribution pattern in coachwood, however, was similar to that found generally in the subtropical rainforest species even though this species grows in differently structured rainforest (Baur, 1965). Comparisons with other species (Table 4) were based on results for mature leaves on the tree and fresh litter using comparable sampling techniques. Fully-developed younger leaves have nutrient concentrations different to those in older leaves, so that there are different patterns of redistribution taking place within the crown. Further, there are different patterns between different-aged forests of the same species (Ashton, 1975). The pattern for nitrogen was similar to that for phosphorus for the rainforest species. Orites excelsa was again found to form a different pattern. Nitrogen was generally not as efficiently redistributed as phosphorus during heartwood formation. Ceratopetalum apetalum and Lophostemon confertus in this case followed the rainforest pattern, while the eucalypts were differently distributed. While for most nutrients there are relationships between foliage litter and heart- wood redistribution which may be described as specific to species/site, phosphorus and nitrogen have given separate patterns of nutrient cycling. In rainforests, where phos- phorus and nitrogen are usually readily available, a certain amount of ‘energy’ is apparently expended in retaining nutrients in biomass and hence not all tissues are affected. In the case of the lower phosphorus sclerophyllous forest, all available phos- phorus is apparently redistributed, this being the primary limiting nutrient. The pattern of Orites excelsa possibly indicated that it evolved in a low phosphorus environment. References ASHTON, D. H., 1976. — Phosphorus in forest ecosystems at Beenak, Victoria. /. Ecol. 64: 171-186. ATTIWILL, P. M., 1980. — Nutrient cycling in a Eucalyptus obliqua (L-Hérit.) forest — IV. Nutrient uptake and nutrient return. Aust. J. Bot. 28: 199-222. PROG. LINN. SOC. N.S.W., 111 (1), 10 NUTRIENT REDISTRIBUTION IN RAINFOREST TREES Baur, G. W., 1957. — Nature and distribution of rain-forests in New South Wales. Aust. J. Bot. 51: 190-222. — , 1965. — Forest types in New South Wales. For. Comm. N.S.W. Res. Note No. 17. BECKMAN, G. G., and THOMPSON, C. H., 1976. — The soils. In The Border Ranges — a land use conflict in regional perspective (cds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Qucensland. BurGess, I. P., FLoyp, A., KikKAwA, J., and PATTIMORE, V., 1975. — Recent developments in the silvi- culture and management of subtropical rainforest in N.S.W. Proc. Ecol. Soc. Aust. 9: 74-84. HINGSTON, F. J., TURTON, A. G., and DIMMOCK, G. M., 1979. — Nutrient distribution in Karri (Eucalyptus diversicolor F. Muell.) ecosystems in southwest Western Australia. For. Ecol. Managem. 2: 133-158. HORNE, R., and GWALTER, J., 1982. — The recovery of rainforest overstorcy following logging. I. Subtropi- cal rainforest. Aust. For. Res. 13: 29-44. JENNY, H., GesseL, S. P., and BINGHAM, F. T., 1949. — Comparative study of decomposition rates of organic matter in temperate and tropical regions. Soz/ Scz. 68: 419-432. KELLY, J., LAMBERT, M. J., and TURNER, J., 1983. — Available phosphorus forms in forest soils and thcir possible ecological significance. Commun. Soil Sci. Plant Anal. 14: 1217-1234. LAMBERT, M. J., 1981. — Inorganic constituents in wood and bark of New South Wales forest tree species. For. Comm. N.S.W., Res. Note 45. 43 pp. —., 1983. — Methods for chemical analysis. For. Comm. N.S.W. Tech. Pap. 25. Third Edition. 187 pp. , and TURNER, J., 1983. — Soil nutrient-vegetation relationships in the Eden area, N.S.W. III. Foliage nutrient relationships with particular reference to Eucalyptus sub genera. Aust. For. 46: 200-209. , and , 1986. — Nutrient concentrations in foliage of species within a New South Wales sub-tropical rainforest. Ann. Bot. 58: 465-478. . , and KELLY, J., 1983. — Nutrient relationships of tree species in a New South Wales sub-tropical rainforest. Aust. For. Res. 13: 91-102. OLSON, J. S., 1963. — Energy storage and the balance of producers and decomposers in ecological systems. Ecology 44: 322-331. ? RICHARDS, B. N., and CHARLEY, J., 1977. — Carbon and nitrogen flux through native forest floors. In Nutrient cycling in indigenous forest ecosystems: 65-81. Perth: C.S.I.R.O. Division of Land Research and Management. STEVENS, N. C., 1976. — Geology and landforms. In The Border Ranges — a land use conflict in regional perspective (eds, R. MONROE and N. C. STEVENS). Brisbane: Royal Society of Queensland. TURNER, J., 1975. — Nutrient cycling in a Douglas-fir ecosystem with respect to age and nutrient status. Seattle, Washington: University of Washington, Ph.D. thesis, unpubl. , and KELLY, J., 1981. Relationships between soil nutrients and vegetation in a north coast forest, New South Wales. Aust. For. Res. 11: 201-208. , and LAMBERT, M. J., 1983. — Nutrient cycling within a 27-year-old Eucalyptus grandis plantation in New South Wales. For. Ecol. Managem. 6: 155-168. WEBB, L. J., 1969. — Edaphic differentiation of some forest types in eastern Australia. II. Soil chemical factors. J. Ecol. 57: 817-830. APPENDIX List of common names, scientific names and authorities of species from the study site Common name Bangalow palm Birds nest fern Botanical name Archontophoenix cunninghamiana (H. Wendl.) H. Wend. et Drude Asplenium nidus L. Black booyong Heritiera actinophylla (F. M. Bail.) Kosterm. Bollygum Neolttsea reticulata (Meisn.) F. Muell. Brushbox Lophostemon confertus (R. Br.) Peter G. Wilson e J. T. Waterhouse Coachwood Ceratopetalum apetalum D. Don Corkwood Caldcluvia paniculosa (F. Muell.) Hoogl. Doughwood Euodtia micrococca F. Muell. Giant stinging tree Dendrocnide excelsa (Wedd.) Chew Kangaroo apple Solanum aviculare Forst. f. Lace fern Nephrolepis sp. Pigeonberry ash Cryptocarya erythroxylon Maiden et Betche Prickly ash Orites excelsa R. Br. Red carabeen Geissois benthamiana F. Muell. Rosewood Dysoxylum fraseranum (A. Juss.) Benth. Teak Flindersia australis R. Br. Tree fern Cyathea australis (R. Br.) Domin White booyong Heritiera trifoliolata (F. Muell.) Kosterm. Wild tobacco tree Yellow carabeen Solanum mauritianum Scop. Sloanea woollsii F. Muell. PROC. LINN. SOC. N.S.W., 111 (1), 1989 New Occurrences of Culmacanthid Acanthodians (Pisces, Devonian) from Antarctica and southeastern Australia G. C. YOUNG YOUNG, G. C. New occurrences of culmacanthid acanthodians (Pisces, Devonian) from Antarctica and southeastern Australia Proc. Linn. Soc. N.S.W. 111 (1), 1989: 11-24. Two new spccies of the acanthodian fish Culmacanthus Long 1983 are described from the lower part of the Aztec Siltstone of southern Victoria Land. Antarctica, and Facies 3 of the Boyd Volcanic Complex of southeastern New South Wales. Both are represented only by cheek plates. That of C. antarctica sp. nov. has distinctive ornament, a longer lateral than dorsal lamina, and the infraorbital sensory groove passing off the ventral margin of the plate. C. pambulensis sp. nov. resembles the type species C. stewarti Long in the shape of the posterior margin of the check plate and the ventral course of the infraorbital sensory canal, but differs in its proportions, the shape of the anterior margin, and the much finer dermal ornament. C. antarctica sp. nov. is considered to be the oldest (?late Middle Devonian) because it is associated with turiniid thelodont scales. The two other species of Culmacanthus occur with bothriolepid and phyllolepid placoderms in assemblages considered to be early Late Devonian (Frasnian). The specific differences described herein suggest however that they are not precise cor- relatives. Culmacanthid acanthodians are only known from southeastern Australia and southern Victoria Land, a distribution pattern previously noted in chondrichthyans and placoderms from the same faunas, and suggesting that Culmacanthus was an cast Gondwanan endemic. G. C. Young, Division of Continental Geology, Bureau of Mineral Resources, PO. Box 378, Canberra, Australia 2601; manuscript received 15 December 1987, accepted for publication 24 August 1988. INTRODUCTION The acanthodians are a major group of Palaeozoic gnathostome fishes, with a fossil record from Silurian to Permian. During the Devonian Period they were widely dis- tributed in both non-marine and marine aquatic environments, and their tiny scales and characteristic fin-spines are commonly represented in microvertebrate assem- blages. However articulated specimens are much less common, and knowledge of acanthodian morphology is based mainly on a few localities in Europe and North America which have yielded well-preserved complete specimens (for a comprehensive review of the group see Denison, 1979). A. S. Woodward was the first to describe Australian acanthodians, from the Lower Carboniferous Mansfield group of Victoria (Woodward, 1906), and he also published the first descriptions of the group from Antarctica (Woodward, 1921). Isolated acantho- dian scales and spines occur widely in Devonian rocks of eastern and central Australia (Fig. 1), and in recent years some well-preserved articulated acanthodians have been described from southeastern Australia (e.g. Long, 1983a, 1986a). The best-known locality is Mount Howitt in east central Victoria, which is the type locality for the genus Culmacanthus Long 1983, of which additional material is described in this paper. With these two new species the genus Culmacanthus is now known from three localities in southeastern Australia, and one locality in Antarctica (Fig. 1). Its distribution pattern matches that seen in several other taxa of Devonian fishes, the palaeogeographic sig- nificance of which is discussed below. All described specimens are housed in the Com- monwealth Palaeontological Collection (prefix CPC), Bureau of Mineral Resources, PROC. LINN. SOC. N.S.W., 111 (1), 1989 12 DEVONIAN FOSSIL FISHES abs Als MR WC 4 A ARS AUSTRALIA EAST ANTARCTICA 20/09/172 Fig. 1. Reconstruction of East Antarctica against Australia, modified from the Gondwana reconstruction of Lawyer and Scotese (1987). showing the main localities of Devonian acanthodian fishes in the southwest Pacific region. Numbered localities are the only known occurrences of culmacanthid acanthodians, as dealt with in this paper; 1, Mount Crean, Lashly Range, southern Victoria Land (Culmacanthus antarctica sp. nov.); 2, Mount Howitt, cast central Victoria (Culmacanthus stewart: Long); 3, Freestone Creek, east Gippsland, (culmacanthid indet.); 4, Pambula River, southeastern New South Wales (Culmacanthus pambulensis sp. nov.). Other abbreviations are: B, Broken River, Queensland; BD, Burrinjuck Dam, New South Wales; C, Cobar area, Darling Basin; DS, Dulcie Syncline, Georgina Basin, EM, Crashsite Quartzite, Ellsworth Mountains; G, Grampians, western Victoria; H, Horlick Formation, Ohio Range; M, Munyarai, Officer Basin; MBL, Marie Byrd Land, West Antarctica, MR, Macdonnell Range, Amadeus Basin; NNZ, New Zealand, North Island; R, Reefton, New Zealand, South Island; RS, Ross River Syncline, Amadeus Basin; T, Tasmania; TS, Toko Syncline, Georgina Basin; WC, Wilson Cliffs, Canning Basin. Geology and Geophysics, Canberra, A.C‘T. Material of the type species Culmacanthus stewarti Long is housed in the Museum of Victoria (prefix NMP). PREVIOUS INVESTIGATIONS The first samples of the Antarctic Aztec fish fauna were collected by F. Debenham on the British Antarctic “Ierra Nova expedition of 1910-13, from moraine material at PROC. LINN. SOG. N.S.W., 111 (1), 1989 G. C. YOUNG 13 Granite Harbour on the coast of southern Victoria Land. They were described by Woodward (1921), who identified eight taxa of Devonian fishes, including isolated acanthodian scales which were said to resemble those of Chezracanthus murchisoni from the Middle Devonian of Scotland. /n situ remains of the Aztec fauna were not discovered until the Trans-Antarctic Expedition of 1955-58 (see Gunn and Warren, 1962), when Devonian fish remains were collected from the Aztec Siltstone at three localities (Lashly Mountains, Mount Feather, Boomerang Range) in the region of the Skelton Névé of southern Victoria Land (locality 1, Fig. 1). Other fish remains from the same general area were recorded by Matz and Hayes (1966) and Matz, Pinet and Hayes (1972; their ‘Fortress Formation’) from west Beacon Heights, Mount Fleming, Mount Crean, and Aztec Mountain, and by Helby and McElroy (1969) from Aztec Mountain. Gunn and Warren's collection was described by White (1968). The most extensive collections of Devonian fishes from the Aztec Siltstone were made by Antarctic expeditions from Vic- toria University of Wellington in 1968-69 (see McKelvey et al., 1972), and in 1970-71 (VUWAE 15), when the specimen described below was collected. Detailed measured sections through the Aztec Siltstone at most of the fossil localities are given in Askin et al. (1971) and Barrett and Webb (1973). Recent publications dealing with or discussing the Aztec fish fauna include Ritchie (1971a, b, 1974, 1975). Young (1982, 1988), and Grande and Eastman (1986). A detailed locality map and a summary of the 24 known localities for the Aztec Siltstone fish fauna are given in Young (1988). In the material collected by Gunn and Warren, White (1968) identified several types of acanthodian spines. Ribbed spines from the upper fossiliferous horizon in the Boomerang Range were referred to a new species (G. warren) of the genus Gyracanthides, originally described by Woodward (1906) from the Early Carboniferous of Mansfield in Victoria. A new genus and species (Antarctonchus glacialis White) was erected for long slender acanthodian spines with numerous strong longitudinal ribs. White also deter- mined as an acanthodian some small spine fragments originally described by Woodward (1921) as an antiarch (Byssacanthoides debenhami Woodward). In addition to these named taxa from the Middle-Late Devonian, V. T. Young (1986) has referred to the genus Machaeracanthus two types of Early Devonian acanthodian spines from the Horlick Formation in the Ohio Range of the southern Transantarctic Mountains, and a similar specimen is recorded from the Ellsworth Mountains (H, EM, Fig. 1). In southeastern Australia acanthodian remains have been known from Victoria for some time (Woodward, 1906; Hills, 1931). More recently Long (1983a) described Culmacanthus stewart: from the Mount Howitt fish locality, and figured one incomplete specimen from Freestone Creek in eastern Victoria as a probable culmacanthid (locali- ties 2, 3, Fig. 1). A second acanthodian from the Mount Howitt fauna (Howzttacanthus) was described by Long (1986a). A summary of the Victorian Devonian fish localities and their biostratigraphy is given in Long (1983b: fig. 1). Placoderms associated with Culmacanthus are also described by Long (1984) and Long and Werdelin (1986). Over the state border, in southeastern New South Wales, Devonian fish remains were discovered in the Eden-Pambula area in 1978 (locality 4, Fig. 1), and a preliminary report (Young, in Fergusson et al., 1979) identified four assemblages ranging in age probably from iatest Middle Devonian (Givetian) through to the end of the Late Devonian (Famennian). One specimen, first described as a possible osteichthyan plate (Fergusson et al., 1979: 103), is referred below to Culmacanthus on the basis of Long’s (1983a) description of this genus. Further collecting in 1979 produced the new speci- mens described below. Other described elements of the Devonian fish faunas from the south coast of New South Wales include sharks from the Bunga Beds near Bermagui (Young, 1982), and an asterolepid antiarch from the Pambula River area (Young, 1983). Elements still to be described include bothriolepid and remigolepid antiarchs, phyllo- PROG. LINN. SOC. N.S.W., 111 (1), 1989 14 DEVONIAN FOSSIL FISHES lepid placoderms, and rhipidistian and other acanthodian remains. A locality map of the region is given in Young (1983: fig. 1). SYSTEMATIC DESCRIPTIONS Subclass ACANTHODII Owen Order CLIMATIIDA Berg Suborder DIPLACANTHOIDEI Miles 1966 Family CULMACANTHIDAE Long 1983 Genus Culmacanthus Long Remarks: The above taxa are defined in Denison (1979) and Long (1983a). Culmacanthus antarctica sp. nov. (Figs. 2A, 3A) 1986a. ‘culmacanthid cheek plates’ (pars) Long, p. 13. Holotype: CPC 26579, a right cheek plate preserved in part and counterpart. Locality: Mount Crean, east side of the Lashly Range (77° 53'S, 159° 33’E), southern Victoria Land, Antarctica (locality 8 of Young, 1988: fig. 3). Horizon: From the lower part of the Aztec Siltstone, within 40 m of the base of the for- mation (unit 5 of section L2 measured by Askin et al., 1971; for a detailed discussion of the stratigraphy of the Mount Crean section see Young, 1988). Diagnosis: A Culmacanthus possessing a cheek plate in which the infraorbital sensory canal passes off the ventral margin, the anteroventral corner is pronounced, the ventral lamina is longer and deeper than the dorsal lamina, and the plate is fairly flat, ornamented with closely spaced ridges and elongate tubercles with a flat external surface, and has a breadth/length index of about 79. Remarks: Long (1983a) did not present a diagnosis for the type species C. stewart, but compared to the Antarctic specimen there are obvious differences in the form of the cheek plate. In the type species the coarser ornament of more widely spaced ridges have a rounded rather than flat dorsal surface, the anterior margin of the plate is distinctly notched (Fig. 3B), there is a more pronounced angle between the dorsal and lateral laminae, the dorsal lamina is longer than the lateral, and the latter has a concave exter- nal surface. The most obvious difference is in the course of the main sensory groove, with both dorsal and ventral sections passing off the anterior margin of the plate. The articulated remains of C. stewart: show that Culmacanthus was a diplacanthoid acanthodian with two dorsal fin-spines, paired pectoral and pelvic spines, and an anal spine (Fig. 5). These spines are ornamented with approximately nine coarse longitudi- nal ribs, and they have a deeply inserted base which is finely striated (Long, 1983a). It is possible therefore that some of the similar acanthodian spines from the Aztec Siltstone named by White (1968) as Antarctonchus may also belong to Culmacanthus. Long (1983a) has already suggested this for the isolated spines from Freestone Creek in Victoria named Striacanthus by Hills (1931). However, in the absence of articulated material demonstrating this, and because the histology of Culmacanthus spines is at present unknown, I follow Long’s (1983a) procedure here, and leave Antarctonchus as a form genus for isolated spines, until such time as new material or more detailed study can demonstrate that the spines and dermal cheek plates come from the same fish. Description: The holotype and only referred specimen 1s an isolated dermal cheek plate preserved in part and counterpart. The external impression was cleaned in dilute hydrochloric acid to remove the bone, and the distinctive ornament is well displayed in a latex cast (Fig. 2A). The counterpart is preserved as bone except for the posterolateral corner, which is an impression of the visceral surface. All margins of the plate are PROC. LINN. SOG. N.S.W., 111 (1), 1989 G. C. YOUNG 15 Fig. 2. Right cheek plates of Culmacanthus in external view (latex casts whitened with ammonium chloride). A, C. antarctica sp. nov., holotype, CPC 26579 (x4). B-D, C. pambulensis sp. nov.; B, holotype, CPC 26580 (x4); C, CPC 26581 (x3); D, CPC 26582 (x4). complete. The orientation of such isolated cheek plates is not readily determined, but following the work of Long (1983a) on the articulated Mount Howitt material of C. stewart, the position of the short sensory groove (ifc.b, Fig. 3A) shows this to be a right dermal cheek plate. It is about 21.5mm long, and relatively deep (breadth/length index of 79). The anterior margin lacks the distinct orbital notches seen in the type species (Fig. 3B), and slopes anteroventrally to a pronounced anteroventral corner, behind which the infraorbital sensory groove passes off the ventral margin of the plate. The plate has a slight flexure at the level of the posterior angle of the infraorbital sensory groove (ifc, Fig. 3), as previously described by Long (1983a), but overall it is a much flatter plate than in the Victorian species (based on NMP 159838). The inflexion of the infraorbital groove is placed high on the plate, which gives it a much larger lateral than dorsal lamina. The short posterodorsal sensory groove (ifc.b), as described for C. stewarti, was interpreted by Long (1983a) as possibly part of the preopercular sensory groove, but this groove in fishes normally passes posterolaterally across the cheek towards the mandibular joint. It is considered more likely that the short groove on the PROG. LINN. SOG. N.S.W., 111 (1), 1989 16 DEVONIAN FOSSIL FISHES cheek plate of Culmacanthus is equivalent to the ‘postero-median branch of the infra- orbital canal’ of Diplacanthus identified by Watson (1937; ‘central sensory line’ of Miles, 1966). Fig. 3. Cheek plates of Culmacanthus restored in external view. A, a right plate of C. antarctica sp. nov., after the holotype, CPC 26579; B, C. stewart: Long, a left plate, restored after a latex cast of NMV P159838 (see Long 1983a: fig. 4B); C, aright plate of C. pambulensis sp. nov., after the holotype, CPC 26580. an, anterior notch; dep.p, posterior depression; ifc, infraorbital sensory groove; ifc.b, posterodorsal (‘central’) branch of infra- orbital sensory groove; pn, posterior notch. The distinctive ornament of C. antarctica sp. nov. comprises cusp-like tubercles which change into elongate ridges towards the posterior margin. In front of the sensory groove the surface of each cusp slopes upwards from the bone surface in a posterodorsal direction, at which a point may be developed, to give some cusps a triangular shape. Behind the sensory groove the more elongate cusps and short ridges have their pointed ends directed posteriorly, and are separated by deep, narrow grooves. Most of the ridges and tubercles are ornamented with up to four very fine ridges which may branch towards the anterior. Typically the elongate primary ridges of the ornament which are most pronounced on the posterior half of the plate have a flat or slightly concave upper surface, with the edges clearly marked by two fine ridges. Compared to C. stewartt (NMP 159838, Fig. 3B) the ornament is more finely sculptured, with the ridges and tubercles more closely spaced. In the type species (Long, 1983a, fig. 4A, B) the ornament is PROC. LINN. SOG. N.S.W,, 111 (1), 1989 G. C. YOUNG Vi) coarser, with wider intervening spaces, and scattered tubercles along the ventral margin. However some fine striations are also visible on the more elongate tubercles and ridges, and in larger specimens there are fewer tubercles and longer ridges (Long, 1983a, fig. 2B). Culmacanthus pambulensis sp. nov. (Figs. 2B-D, 3C, 4) 1979. ‘osteichthyan, with an ornament of delicate subparallel ridges’ Young, in Fergus- son et al., p. 103. 1983a. ‘dermal cheek plate. . . from Pambula River’ Long, p. 54. 1986a. ‘culmacanthid cheek plates’ (pars) Long, p. 13. Fig. 4. Culmacanthus pambulensis sp. nov., cheek plates in external view. A, a right plate, after CPC 26581; B, a left plate, after CPC 26583. For abbreviations see caption to Fig. 3. PROC. LINN. SOG. N.S.W., 111 (1), 1989 18 DEVONIAN FOSSIL FISHES Name: Abbreviated from the nearby town of Pambula, N.S.W. Holotype: CPC 26580, a right cheek plate. Other Material: Right (CPC 26581, 26582) and left (CPC 26583) cheek plates, and another ornamented fragment (CPC 17004). Locality: Grid Reference 7510E 59081N on the Pambula 1:25000 Sheet, about 5km west of the town of Pambula on the south coast of New South Wales (see Young, 1983: fig. 1). Horizon: Facies 3 of the Boyd Volcanic Complex, as described by Fergusson et al. (1979). Diagnosis: A Culmacanthus possessing a fairly flat cheek plate in which the infraorbital sensory canal passes off the anterior margin, the anterior margin has only shallow or in- distinct notches, the ventral lamina 1s considerably deeper than the dorsal lamina, the closely spaced ornament comprises fine ridges and elongate tubercles, and the breadth/length index varies between at least 68 and 94. Remarks: Apart from the obvious and consistent differences in ornament, the three measurable specimens of C. pambulensis are broader than in the type species, although for the smallest example described below (CPC 26580) this difference is only slight. Long (1983a: 56) regarded such differences in proportion to be generic rather than specific features, but the variability in proportions is clear in this new species, and the other obvious similarities indicate that these specimens are properly referred to the genus. C. pambulensis resembles C. antarctica rather than C. stewarti in the flatter cheek plate, the longer and steeper disposition of the ventral section of the infraorbital sensory groove, and the absence or slight development of notches on the anterior margin. It differs from C. antarctica but resembles C. stewarti in the fact that the infraorbital groove passes off the anterior margin of the plate. Long (1983a) noted other specimens from Freestone Creek in eastern Victoria with broader proportions than the type species, as in C. pambulensis sp. nov., but that material is at present too poorly known to permit useful comparisons. Description: All specimens are preserved as external moulds, which have been studied by latex casting. The holotype (Figs 2B, 3C) is the smallest specimen (length 14mm), and the most complete. The infraorbital groove has an attenuated ventral section, not seen in other examples, and interpreted as an individual variation. The posterodorsal branch is a short separate groove (ifc.b, Fig. 3C) as in other examples with this part preserved. Compared to C. stewart: (represented by NMP 159838) the cheek plate is rather flat. The lateral lamina is slightly concave near the posterior corner (dep.p), and there may be another shallow depression running anteroventrally beneath the ventral section of the infraorbital canal, but otherwise both laminae of the plate are gently con- vex. The posterior depression is seen in all available specimens of this species (Fig. 2B- D). Two larger specimens are unusually broad (Fig. 4), although their posterior margins are somewhat irregular. This may be due to incomplete preservation, but an irregular margin abutting against small plates at the posterior margin of the gill cover is con- sidered more likely. It is possible that breadth of the plate increased allometrically with size, but evidence against this is provided by C. stewartz, where the largest plate (Long, 1983a: fig. 2) is also the most elongate (erroneously stated as ‘twice as broad as long’ by Long, 1983a: 56). In both CPC 26581 and 26583 the lateral lamina projects posteriorly, as in C\ antarctica described above, and all specimens with the region preserved show a shallow notch on the posterodorsal margin (pn, Figs 3C, 4). The general similarity of the delicate ornament in the two largest examples (Figs 2C, 4B) to other specimens from this locality suggests that they are conspecific, and that proportions of the cheek plate are more variable than previously thought. In the type species only three cheek plates have been described, with two from one individual, so further material is required to assess the variability of this feature. PROC. LINN. SOC. N.S.W., 111 (1), 1989 G. C. YOUNG 19 The ornament in C. pambulensis is much finer than in C. antarctica or C. stewarti. Small triangular tubercles with posterodorsally directed points predominate in front of the sensory canal, as in the other species, and behind the canal these tubercles become more elongate and posteriorly directed (Fig. 2B, D), and may coalesce into subparallel ridges (Figs 2C, 4B). Specimen CPC 17004 previously mentioned by Young (in Fergus- son et al., 1979) is referred to C. pambulensis only on its similar ornament, since no sen- sory grooves are preserved. DISCUSSION Morphology and Systematics Denison (1979) included two genera in his family Diplacanthidae — Dizplacanthus Agassiz and Gladiobranchus Bernacsek and Dineley. Long (1983a) preferred to place Gladiobranchus as closely related to Uraniacanthus, as originally suggested by Bernacsek and Dineley (1977). However Bernacsek and Dineley supported this relationship by suggesting that Uraniacanthus may have possessed dermal plates and prepectoral spines, even though Miles (1973) expressed no doubt from detailed study of the specimens that these structures were absent in Uranzacanthus. Bernacsek and Dineley (1977) also inferred that Gladiobranchus had dentigerous jawbones, but the holotype shows anterior circum- orbital bones, indicating proximity of the snout, but no sign of ossified jaw elements. These may have been lost, but the available evidence favours the interpretation that Gladiobranchus did not possess them. Supporting this, and the climatiid affinities of Gladiobranchus, are the presence of dermal shoulder plates, prepectoral spines, and the large plates and tesserae of the head (Denison, 1979). The scapulocoracoid of Gladio- branchus was restored by Bernacsek and Dineley (1977) as alow and broad element with a truncated dorsal end, but the specimen (Bernacsek and Dineley, 1977: pl. 7) suggests a higher narrow dorsal termination, with an anterior ridge and expanded posterior ventral part, just as in Diplacanthus. The two enlarged circumorbitals, one at the posterodorsal corner of the orbit, are special features shared with Dzplacanthus, as Denison (1979) noted. Thus I follow Denison (1979) in considering Gladiobranchus better placed as a diplacanthoid acanthodian, rather than an ischnacanthid (see also Long, 1986b: 335). Long (1983a) modified Miles’s (1966) diagnosis of the Diplacanthoidei to include Culmacanthus. The most significant resemblances (Fig. 5) are in the large dermal cheek plate crossed by the infraorbital sensory groove, the relatively deep body, and the deeply inserted spines. Culmacanthus differs from Diplacanthus primarily in the larger size of the cheek plate, the absence of intermediate fin-spines, the probable absence of a mandi- bular bone, the high narrow scapulocoracoid, and the structure of the dermal shoulder girdle, with an unpaired lorical plate, and no spines attached to the paired pinnals. Long (1983a) suggested that the cheek plate of Culmacanthus may have been homologous to the postorbital or preopercular bones of Diplacanthus, but since there is no sensory groove passing posterolaterally across the plate it seems more likely that it is only a much en- larged postorbital, which extended more dorsally than the corresponding bone of Diplacanthus, to include the junction of the posterodorsal branch of the infraorbital canal at its ossification centre. For functional reasons it is likely that the cheek plate only covered the region of the palatoquadrate, with a flexible hyoid gill cover situated between it and the scapulocoracoid. The well-ossified branchiostegal rays in this position in Gladiobranchus may be interpreted as a symplesiomorphy, by outgroup com- parison with climatiids. It is unclear from Long’s (1986b) analysis of the dermal shoulder girdle of climatiids whether the condition in Culmacanthus is primitive or specialized, but if intermediate fin-spines and pectoral spines firmly attached to the pinnal plates are PROC. LINN. SOC. N.S.W., 111 (1), 1989 20 DEVONIAN FOSSIL FISHES Fig. 5. Culmacanthus stewartt Long. Restoration of complete fish, slightly modified after Long (1983a: fig. 9). synapomorphies of ‘higher climatiids’ as he suggests (1986b: 337), then Dzplacanthus would be more closely related to some Early Devonian climatiids than to Culmacanthus. All students of acanthodian evolution have acknowledged the small number of reliable characters available to define the major groups, and in the case of Culmacanthus the main evidence still to be forthcoming concerns the histology of the scales and fin spines. This is not known in the Victorian material because the bone is badly weathered. Better- preserved material permitting histological study can be expected to clarify the relation- ships of this form to the Euramerican diplacanthids. Buostratigraphy A biostratigraphic analysis of the Aztec fish fauna based on the distribution of antiarchs was presented by Young (1988), in an attempt to resolve previous differences of opinion about its precise age. Woodward (1921) first proposed a Late Devonian age on the evidence of the antiarch Bothriolepis, and associated shark and large osteolepid scales, and Gross (1950) reached the same conclusion. However White (1968) noted the apparent mixing in the Aztec fauna of groups that elsewhere range in age from Middle or even Early Devonian, to Late Devonian or Early Carboniferous, and on this basis he favoured a late Middle Devonian rather than early Late Devonian age. Ritchie (1975) added the placoderms Phyllolepis and Groenlandaspis to the faunal list, and proposed a younger Late Devonian age. Young (1982) suggested correlations with southeastern Australia based on the sharks in the Aztec fauna, which were consistent with Helby and McElroy’s (1969) assessment of a Frasnian age for a palynoflora dominated by the spore Geminospora lemurata. This species was assessed by Playford (1983) to have a known bio- stratigraphic range from early or middle Givetian to late Frasnian or early Famennian. However in Europe McGregor (1979) has estimated a much greater (Emsian- Famennian) range for the possibly synonymous Gemuinospora svalbardiae, so the spore evidence at present only dates the Aztec Siltstone within broad limits. Other evidence supporting comparisons between the Aztec succession and the Devonian sequences of southeastern Australia include the acanthodian Gyracanthides described from Victoria by Woodward (1906), and from the upper Aztec succession by White (1968), the antiarch Pambulaspis from southeastern New South Wales (Young, PROC. LINN. SOC. N.S.W., 111 (1), 1989 G. C. YOUNG 21 1983) and described by Young (1988) from the top of the Aztec, and the culmacanthid acanthodians just described. Culmacanthus antarctica 1s associated with turiniid thelodont scales and bothriolepid antiarchs, and may therefore be the oldest known species, since no thelodonts have been recorded from the successions in eastern Victoria and the south coast of New South Wales. Young (1988) noted that the association of bothriolepid anti- archs with thelodont scales is currently known from only one other locality, in the Hatchery Creek fauna from near Wee Jasper, New South Wales (Young and Gorter, 1981). A post-Emsian age for this fauna is indicated by conodonts in the underlying Murrumbidgee Group (serotznus Zone or slightly younger), but a younger age limit is un- certain. In contrast the two Australian species of Culmacanthus are associated with phyllolepid placoderms, which occur widely in southeastern Australia, in all other post- Emsian faunas currently known which contain placoderms. From eastern Victoria and the southern coast of New South Wales only the Bunga Beds fauna (Young, 1982) lacks phyllolepids, but this is an impoverished fauna in which placoderms have not been found. The abundant placoderms in the Aztec Siltstone includes phyllolepids only in the highest horizons, and Young (1988) suggested that the base of the phyllolepid zone preserved in the Aztec Siltstone, and the first appearance of the antiarch Pambulaspis, may be older than the occurrence of related forms in eastern Victoria and southeastern New South Wales. In eastern Victoria, Long (1983b: fig. 3) used the appearance of Bothriolepis and phyllolepids to define the base of the Upper Devonian, with Groenlandaspis occurring in the latter half of the Frasnian. The fish fauna in the Boyd Volcanic Complex (Fergusson et al., 1979; Young, 1982, 1983), which as well as Culmacanthus includes at least four taxa (Bothriolepis, Pambulaspis, Antarctilamna, phyllolepids) in common with the Aztec fauna, apparently lacks Groenlandaspis. The fact that both Bothriolepis and Groenlandaspis in the Aztec Siltstone occur in the lowest fossiliferous beds, and far below the phyllolepid zone, indicates again that these are earlier occurrences than the range of these forms in southeastern Australia. In western New South Wales and central Australia groen- landaspid arthrodires are associated with thelodont scales in the Wuttagoonaspis fauna (Ritchie, 1975; Turner et a/., 1981) but the only antiarchs from this fauna are asterolepids (Young, 1984a), so this appears to be older than the Aztec fauna. An Early-Middle Devonian age has been assigned to the Wuttagoonaspis fauna, but the occurrence of the Antarctic shark Mcmurdodus (Turner and Young, 1987), otherwise only known from the Aztec Siltstone, may indicate less difference in age to the lower Aztec fauna than previously thought. In southeastern New South Wales the Boyd Volcanic Complex provides evidence for a minimum age for the Aztec fauna, because the overlying Merrimbula Group con- tains a marine invertebrate fauna of probable late Frasnian age (e.g. Roberts et al., 1972). A single early Late Devonian marine transgression in southeastern Australia may be assumed, for which the oldest reliable age is late Frasnian based on upper gigas Zone conodonts (Pickett, 1972). This implies a late Givetian — early Frasnian age for the underlying fishes and equivalent faunas in eastern Victoria, and an approximate younger limit to the age of the Aztec Siltstone based on the phyllolepid placoderms in its upper horizons. However, although the Victorian and southern New South Wales fish successions are broadly equivalent in age, detailed taxonomic work on placoderms common to the two areas (e.g. bothriolepids, phyllolepids) has not yet been done. The results of the present study show specific differences between the Culmacanthus remains from the two sequences, which may provisionally be regarded as indicating difference in age, but there is as yet no reliable evidence on which occurrence 1s the older. It should also be noted that according to Long (1983a) the culmacanthid from Freestone Creek is not conspecific with C. stewartz, although other species (e.g. Bothriolepis cullodenensis Long, PROC. LINN. SOG. N.S.W., 111 (1), 1989 22 DEVONIAN FOSSIL FISHES 1983b) are shared between the two assemblages. Further detailed taxonomic work on the placoderms in the various faunas from southeastern New South Wales and Victoria should clarify the precise correlation between the two sequences. Biogeography The biogeography of the Aztec fish fauna has been discussed previously by Young (1981, 1982, 1984b, 1988) and Grande and Eastman (1986), whilst Long (1986a) has recently commented on acanthodian biogeography in the Devonian. The presence of species of Culmacanthus in southeastern Australia and southern Victoria Land, Antarc- tica, but nowhere else, corresponds with the pattern indicated previously by the elasmo- branch Antarctilamna (Young, 1982), the antiarch Pambulaspis (Young, 1988), and possibly the placoderm Antarctaspis (if a relationship to phyllolepids is confirmed; see Young, 1987: fig. 5). These distributions are all consistent with grouping the two regions together in an ‘East Gondwana Province’ as proposed by Young (1981). It should also be noted that the only two fish assemblages recorded from any continent in which bothrio- lepid antiarchs and turiniid thelodonts are associated are in southeastern Australia and southern Victoria Land, and species group interrelationships for the cosmopolitan anti- arch Bothriolepis again indicate an Antarctica-Australia connection (Young, 1988). All these fishes inhabited freshwater environments, but this does not mean that they were unable to cross marine barriers, and in this respect no specific data are provided on the Devonian palaeogeography of the Antarctic-Australian region. It is relevant to note however that the distinctive dermal cheek plates of Culmacanthus have not been found in the well-studied fish faunas from the classic Old Red Sandstone localities of the North- ern Hemisphere, but that they do occur in the Southern Hemisphere in two areas separated today by the vast expanse of the Southern Ocean, and a distance in excess of 4500km, which exceeds by a considerable margin the 3000km-wide ocean postulated by some authors to have separated Gondwana from Euramerica during the Late Devonian. Thus the evidence of culmacanthid distribution is entirely consistent with reconstruc- tions of a Palaeozoic Gondwana which minimize or remove oceanic barriers by juxta- posing southeastern Australia and Victoria Land (Fig. 1). ACKNOWLEDGEMENTS The opportunity to participate in the Victoria University of Wellington Antarctic Expedition of 1970-71, and the field and logistic support given to fossil collection in southern Victoria Land, is gratefully acknowledged. R. W. Brown assisted in field collection in the Eden-Pambula region, and took the photographs. W. Peters prepared the specimens, and Dr John Long provided latex casts of Culmacanthus stewart: for com- parison, and commented on the new specimens and the manuscript. Dr R. Ryburn assisted with the bibliography. The paper is published with the permission of the Director, Bureau of Mineral Resources, Canberra. References ASKIN, R. A., BARRETT, P. J., KOHN, B. P., and MCPHERSON, J. G., 1971. — Stratigraphic sections of the Beacon Supergroup (Devonian and older(?) to Jurassic) in South Victoria Land. Publ. Geol. Dept, Victoria University, Antarctic Data Ser. 2: 1-88. BARRETT, P. J., and Wess, P. N., 1973. — Stratigraphic sections of the Beacon Supergroup (Devonian and older(?) to Jurassic) in South Victoria Land. Publ. Geol. Dept, Victoria University, Antarctic Data Ser. 3: 1-165. BERNACSEK, G. M., and DINELEY, D. L., 1977. — New acanthodians from the Delorme Formation (Lower Devonian) of N.W.T., Canada. Palaeontographica A158: 1-25. DENISON, R. H., 1979. — Acanthodii. Handbook of Paleoichthyology. Volume 5. H-P. SCHULYZE (ed.). Stuttgart and New York: Gustav Fischer Verlag. PROC. LINN. SOC. N.S.W., 111 (1), 1989 G. GC. YOUNG 23 FERGUSSON, C. L., CAS, R. A., COLLINS, W. J., CRAIG, G. Y., CROOK, K. A. W., POWELL, C. MCA., Scort, P. A., and YOUNG, G. C., 1979. — The Late Devonian Boyd Volcanic Complex, Eden, N.S.W. Jour. Geol. Soc. Aust. 26: 87-105. GRANDE, L., and EASTMAN, J. T., 1986. — A review of the Antarctic ichthyofauna in the light of new fossil discoveries. Palaeontology 29: 113-137. Gross, W., 1950. — Dic palaeontologische und stratigraphische Bedeutung der Wirbeltierfaunen des Old Reds und der marinen altpalaozoischen Schichten. Abh. dt. Akad. Wiss. Berl. Math. Nat. Kl. 1949: 1-130. GUNN, B. M., and WARREN, G., 1962. — Geology of Victoria Land between the Mawson and Mulock Glaciers. Antarctica. Sczent. Rep. transantarct. Expedition 11: 1-157. He.py, R. J., and MCELRoy, C. T., 1969. — Microfloras from the Devonian and Triassic of the Beacon Group, Antarctica. N.Z. J. Geol. Geophys. 12: 376-382. -HIiuts, E. S., 1931. — The Upper Devonian fishes of Victoria, Australia, and their bearing on the strati- graphy of the state. Geol. Mag. 68: 206-231. Lawver, L. A., and Scoresr, C. R., 1987. — A revised reconstruction of Gondwanaland. In G. D. MCKENZIE (ed.), Gondwana Six: Structure, Tectonics, and Geophysics. Geophysical Monograph 40: 17-23. American Geophysical Union. LONG, J. A., 1983a. — A new diplacanthoid acanthodian from the Late Devonian of Victoria. Mem. Ass. Australas. Palaeont. 1: 51-65. ——, 1983b. — New bothriolepid fish from the Late Devonian of Victoria, Australia. Palaeontology 26: 295-320. ——, 1984. — New phyllolepids from Victoria, and the relationships of the group. Proc. Linn. Soc. N.S.W. 107: 263-308. — , 1986a. — A new Late Devonian acanthodian fish from Mt. Howitt, Victoria, Australia, with remarks on acanthodian biogeography. Proc. Roy. Soc. Vict. 98: 1-17. ——, 1986b. — New ischnacanthid acanthodians from the Early Devonian of Australia, with comments on acanthodian interrelationships. Zool. J. Linn. Soc. Lond. 87: 321-339. , and WERDELIN, L., 1986. — A new Late Devonian bothriolepid (Placodermi, Antiarcha) from Victoria, with descriptions of other species from the state. Alcheringa 10: 355-399. Matz, D. B., and Hayes, M. O., 1966. — Sedimentary petrology of Beacon sediments. Antarct. J. U.S. 1: US;4b=I13}5)- ——.,, Pinrr, P. R., and Hayrs, M. O., 1972. — Stratigraphy and petrology of the Beacon Supergroup, southern Victoria Land. In Antarctic Geology and Geophysics, R. J. ADIE (ed.): 353-358. Oslo: Universitetsforlaget. McGReEGor, D. C., 1979. — Spores in Devonian stratigraphical correlation. Spec. Pap. Palaeontol. 23: 163-184. McKe Lvey, B. C., Wess, P. N., GorTON, M. P., and KOHN, B. P., 1972. — Stratigraphy of the Beacon Supergroup between the Olympus and Boomerang Ranges, Victoria Land. In Antarctic Geology and Geophysics, R. J. ADIE (ed.): 345-352. Oslo: Universitets forlaget. MILEs, R. S., 1966. — The acanthodian fishes of the Devonian Plattenkalk of the Paffrath trough in the Rhineland. Ark. Zool. 18: 147-194. —, 1973. — Articulated acanthodian fishes from the Old Red sandstone of England, with a review of the structure and evolution of the acanthodian shoulder-girdle. Brit. Mus. (Nat. Host.) Bull. (Geol.) 24: 115-213. PICKETT, J., 1972. — Late Devonian (Frasnian) conodonts from Ettrema, New South Wales. /. Proc. Roy. Soc. N.S.W. 105: 31-37. PLAYFORD, G., 1983. — The Devonian miospore genus Geminospora Balme, 1962: a reappraisal based on topo- typic G. lemurata (type species). Mem. Ass. Australas. Paleont. 1: 311-325. RitcHik, A., 1971a. — Ancient animals of Antarctica — Part 2. Hemisphere 15 (12): 12-17. — , 1971b. — Fossil fish discoveries in Antarctica. Aust. Nat. Hist. 17: 65-71. —., 1974. — From Greenland’s icy mountains. Aust. Nat. Hist. 18, 28-35. ——.,, 1975. — Groenlandaspis in Antarctica, Australia and Europe. Nature 254: 569-73. ROBERTS, J:, JONES, P) J., JELL, J. S|, JENKINS, I. B: H., MARSDEN, M. A. H., MeKELLAR, R. G., McKELveY, B. C., and SEDDON, G., 1972. — Correlation of the Upper Devonian rocks of Australia. J. geol. Soc. Aust. 18: 467-490. TURNER, S., JONES, P. J., and DRaPER, J. J., 1981. — Early Devonian thelodonts (Agnatha) from the Toko Syncline, western Oueensland, and a review of other Australian discoveries. BMR J. Aust. Geol. Geophys. 6: 51-69. , and YOUNG, G. C., 1987. — Shark teeth from the Early-Middle Devonian Cravens Peak Beds, Georgina Basin, Queensland. Alcheringa 11: 233-244. Watson, D. M. S., 1937. — The acanthodian fishes. Phil. Trans. Roy. Soc. Lond. 228: 49-146. White, E. I., 1968. — Devonian fishes of the Mawson-Mulock area, Victoria Land, Antarctica. Scient. Rep. transantarct. Exped. 16: 1-26. PROG. LINN. SOG. N.S.W., 111 (1), 1989 24 DEVONIAN FOSSIL FISHES WOODWARD, A. S., 1906. — On a Carboniferous fish fauna from the Mansfield district, Victoria. Mem. National Mus., Melbourne 1: 1-32. ——, 1921. — Fish-remains from the Upper Old Red Sandstone of Granite Harbour, Antarctica. British Antarctic (“lérra Nova’) Expedition, 1910. Nat. Hist. Rep. (Geol. ) 1: 51-62. YOUNG, G. C., 1981. — Biogeography of Devonian vertebrates. Alcheringa 5: 225-243. ——, 1982. — Devonian sharks from south-eastern Australia and Antarctica. Palaeontology 25: 817-843. — , 1983. — A new antiarchan fish (Placoderm1) from the Late Devonian of southeastern Australia. BMR J. Aust. Geol. Geophys. 8: 71-81. ——, 1984a. — An asterolepidoid antiarch (placoderm fish) from the Early Devonian of the Georgina Basin, central Australia. Alcheringa 8: 65-80. ——, 1984b. — Comments on the phylogeny and biogeography of antiarchs (Devonian placoderm fishes), and the usc of fossils in biogeography. Proc. Linn. Soc. N.S.W. 107: 443-473. —, 1987. — Devonian palaeontological data and the Armorica problem. Palaeogeog. Palaeoecol. Palaeoclimat. 60: 283-304. ——, 1988. — Antiarchs (placoderm fishes) from the Devonian Aztec Siltstone, southern Victoria Land, Antarctica. Palaeontographica A202: 1-125. , and Gorter, J. D., 1981. — A new fish fauna of Middle Devonian age from the Taemas/Wee Jasper region of New South Wales. Bur. of Min. Res. Geol. Geophys. Bull. 209: 83-147. YOUNG, V. T., 1986. — Early Devonian fish material from the Horlick Formation, Ohio Range Antarctica. Alcheringa 10: 35-44 PROC. LINN. SOC. N.S.W., 111 (1), 1989 A new Australian Species of Argyrodes Simon (Araneoidea: ‘Theridiidae) which preys on its Host M. R. GRAY and G. J. ANDERSON Gray, M. R., & ANDERSON, G. J. A new Australian species of Argyrodes Simon (Araneoidea: Theridiidae) which preys on its host. Proc. Linn. Soc. N.S.W. 111 (1), 1989: 25-30. A new species of the web-parasite spider genus A7gyrodes is described. This spider has a close association with its only known and larger host, the tent web spider Achaearanea mundula. It is a predator of both the host and its eggs and young and makes use of the host’s leaf retreat as a shelter for its egg sac. M. R. Gray, The Australian Museum, PO. Box A285, Sydney South, Australia 2000, and G. J. Anderson, Dept of Medicine, University of Queensland, Royal Brisbane Hospital, Herston, Aus- tralia 4029; manuscript received 5 January 1988, accepted for publication 19 October 1988. INTRODUCTION Large, complex or communal spider webs may harbour a variety of invertebrates as temporary or semi-permanent residents. The small spiders of the worldwide genus Argyrodes Simon (Exline and Levi, 1962) are common residents in the webs of araneoid spiders. Most live either as commensals which take small prey ignored by the host spider or kleptoparasites which steal prey captured by the host (Robinson and Robinson, 1973; Vollrath, 1979; Whitehouse, 1986). The host spider is often much larger (e.g. Nephila or Argiope spp.) than the resident Argyrodes so that competition for food resources would not seem likely. Several studies have shown, however, that prey stealing can have such an impact that some hosts abandon their webs and build elsewhere (Robinson and Robin- son, 1976; Vollrath, 1981; Rypstra, 1981). A few temperate zone species of Argyrodes have become predators of both their hosts and spiderlings (Kaston, 1965). Trail (1980) has observed host predation by A. fictzlium (Hentz) and A. baboquivari Exline and Levi upon linyphiid host spiders including Frontinella sp., and the uloborid Philliponella owent Chamberlin. A. trigonum (Hentz) is a predator of the araneids Metepeira labyrintha Hentz and Mecynogea lemniscata Walckenaer and the linyphiid Nerene radiata Walckenaer (Wise, 1982; Larcher and Wise, 1985). Whitehouse (1986) noted predation by A. antipodiana O.P. — Cambridge upon its araneid host Avaneus pustulosa Walckenaer while the host was moulting and virtually defenceless. Gray (1983) briefly reported upon a distinctive Argyrodes species from eastern Australia which showed host predation behaviour. A description of this spider and some behavioural observations are given below. Argyrodes incursus new species Figs 4-12 Types: Holotype male (KS 18359), Blackbutt Reserve, Newcastle, New South Wales, March 1986, G. Anderson. Paratypes: Allotype female (KS 18360), 3 females (KS 18361), data as above; 2 females (KS 12254-55), Upper Causeway and Couranga Track, Royal National Park, near Sydney, New South Wales, 4.3.1983, M. Gray and C. Horse- man. Australian Museum coll. Material examined: 2 females, Station 32, southwest of North Hummock, Lord Howe Island, New South Wales, 6.2.1971, M. Gray. PROC. LINN. SOC. N.S.W., 111 (1), 1989 26 HOST-PREDATING ARGYRODES Figs 1-3. 1, Argyrodes incursus, ‘engorged’ female. 2-3, Achaearanea mundula: 2, female; 3, web. Scale lines: Figs 1- 2, 1.0mm. Diagnosis. Male palp with marginal coiled embolus, abdomen with dorsal scute. Female with orange-red spot on posterodorsal abdomen, abdomen rounded. Male. Total length 2.18. Carapace length 0.90, width 0.63. Abdomen length 1.27, width 0.89. Leg 1 segment lengths: femur 1.07, patella 0.33, tibia 0.63, metatarsus 0.65, tarsus 0.45. Carapace, sternum and mouthparts dark brown. Abdomen dark brown dorsally and basally; remainder black apart from 4 small orange spots placed around the base of the spinnerets (Figs 4, 5). Femora and tibiae of legs 1 and 2 dark brown, femora 2 lighter PROC. LINN. SOG. N.S.W., 111 (1), 1989 M. R. GRAY ANDG. J. ANDERSON 27 ventrally; remaining segments reddish brown. Legs 3 and 4 reddish brown, femora lightest. Cheliceral groove with 3 promarginal, 1 retromarginal teeth. A.M.E. largest, other eyes subequal. A.M.E. separated from each other by almost 2 diameters and from A.L.E. by 0.5 diameters; P.M.E. separated by 1.75 diameters. Clypeus with wide, trans- verse groove with numerous anterodorsally directed setae in and below groove. Trans- verse stridulatory ridges run across the posterolateral carapace lobes. Abdomen ovoid, widest behind middle and covered with a dorsal scute which extends anteriorly around the pedicel. A separate scute surrounds the spinnerets (Fig. 5). Dorsal pedicel margin strongly sclerotised with evenly spaced enlarged hair bases as stridulatory picks. Male palp (Fig. 11) with tightly coiled embolus (3 turns) surrounding a raised central tegular area. Embolus supported marginally by a membranous conductor. Radix apical, tri- angular and lying adjacent to the elongate median apophysis. Female. Total length 3.36-4.33. Carapace length 1.29-1.35, width 0.88-0.92. Abdomen length 1.97-3.00, width 1.80-2.55. Leg 1 segment lengths: femur 1.20-1.25, patella 0.35- ‘0.38, tibia 0.80-0.96, metatarsus 0.80-0.88, tarsus 0.50-0.62. Carapace, sternum and mouthparts dark brown. Legs 1-3 dark brown except metatarsi and tarsi which are reddish brown; leg 4 reddish brown. Abdomen ovoid to spherical, black with two in- distinct rows of unpigmented spots anterodorsally. Posterodorsal abdomen with a large orange-red spot (occasionally divided into two unequal spots), bounded anteriorly and posteriorly by 1 or 2 narrow transverse orange bars (Figs 7, 8, 9). Four small irregular orange spots are grouped around the spinnerets. Epigynum (Fig. 6) a low mound sur- mounted by two ovoid fossae separated by about half a diameter. Internal genitalia (Fig. 10) with more or less spherical receptacula separated by about half their width. Sperm ducts strongly convoluted. + MORPHOLOGY AND BEHAVIOUR Both the shape and coloration of the abdomen of Argyrodes incursus are unusual. Elongation, shoulder or posterior tubercles and silvery ornamentation are common features of the Argyrodes abdomen (Exline and Levi, 1962). In unfed female specimens of A. incursus the abdomen is ovoid but expands considerably after feeding into a smooth sphere (Fig. 1). A prominent orange-red spot (absent in males) on the posterodorsal abdomen contrasts markedly with the shiny black coloration of the remainder. The function of this distinctive female colour pattern is not understood but may have some aposematic benefit, such as providing protection from other web-invading predators (e.g. mimetid spiders or wasps). A dark brown dorsal scute covers the male abdomen. The male palp is unusual in that the tegulum forms a central ‘drum’ around which the coiled embolus is wound. A. incursus was always found in association with the ‘tent web’ spider, Achaearanea mundula (LL. Koch). This spider (Fig. 2) is common in open forest habitats in eastern New South Wales and Queensland. Populations with which A. zncursus was associated were located at Newcastle, Sydney, and Lord Howe Island, New South Wales. Ach. mundula builds a complex web consisting of a knockdown maze of threads above a finely woven horizontal sheet (Fig. 3). The curled leaf or litter retreat is placed at the centre of the knockdown maze; this provides a refuge for the spider and its egg sacs (up to 6). The webs are usually built among low understorey foliage. They also harbour commensal/ kleptoparasitic species of Argyrodes plus uloborid and mimetid spiders; the latter are temporary residents, well known as predators of araneoid spiders. Specimens of A. incursus were found singly inside the curled leaf retreats of Ach. mundula individuals during February and March. In the Newcastle population, however, the catching parts of 3 host webs harboured 8, 2 and 2 A. incursus individuals respec- PROG. LINN. SOG. N.S.W., 111 (1), 1989 28 HOST-PREDATING ARGYRODES > 3; EX Figs 4-12. Argyrodes incursus. 4-5, male body: 4, dorsal; 5, lateral. 6, epigynum. 7-9, female abdomen: 7, dorsal; 8, lateral; 9, ventral. 10, female internal genitalia, dorsal. 11, male palp, ventral. 12, upper part of curled leaf retreat of Ach. mundula split open to show A. incursus egg sac near apex. Scale lines: Figs 4-11, 0.5mm; Fig. 12, 3.5mm. PROC. LINN. SOC. N.SW., 111 (1), 1989 M. R. GRAY ANDG. J. ANDERSON 29 tively, with the female hosts still occupying their retreats. Six observations were made of A. incursus females actually feeding upon the carcass of their host spider and/or its eggs or spiderlings, always within the leaf retreat. The dead host spiders were partially silk wrapped and their loosely woven egg sacs, if attacked, were torn open. In the Newcastle population A. zncursus individuals also were found living inside the leaf retreats of empty Ach. mundula webs; in one case a male and female were found together. Some of these females had built single egg sacs at the leaf retreat apices (Fig. 12). Seven egg sacs were observed, all with the characteristic urn shape found in many species of Argyrodes (Exline and Levi, 1962). When present, the female sat above its sac on the stalk attachment. At present Ach. mundula is the only known prey of A. zncursus. The capacity for almost tick-like engorgement of the abdomen of A. incursus after feeding is particularly striking. Females may be able to survive and reproduce on the food reserves obtained from one successful predatory episode. The limited observations suggest that mating may occur within the Ach. mundula retreat. In addition, A. incursus females use the empty host leaf retreats for the protected placement of their egg sacs. All of this suggests a close, perhaps exclusive, association of A. zncursus with its host/prey species, Ach. mundula. In a study of A. trigonum (Hentz), Larcher and Wise (1985) observed a wide range of behaviour including commensalism, kleptoparasitism, web theft and host predation. They concluded that the nature of the interaction between A. trigonum and its hosts varied as a function of their relative individual sizes. This agrees with an earlier finding of Trail (1980) that Argyrodes species which attack their hosts are equal or larger in adult size compared with their host species. Presumably, this makes the attacker better able to Overpower its prey. A. incursus, however is considerably smaller than its prey, Ach. mundula. Carapace length ranges for females of A. zncursus and Ach. mundula are respec- tively 1.2971.35mm (M = 1.32) and 2.17-2.58mm (M = 2.32). The mean difference of 1.00mm is highly significant (P< 0.001). The suggestion of Trail (1980) that an Argyrodes species smaller than its host would behave as a commensal or kleptoparasite rather than as a host predator is not supported in this case. As yet, little information about the attack behaviour of A. incursus is available. It seems likely that the host is attacked in its retreat as all carcasses were found in retreats. Limited observations by Larcher and Wise (1985) suggest that A. trigonum simply approaches the host and bites it on a leg, resistance being negligible (though some hosts flee to avoid an attack). The venom is apparently quick to immobilise the prey which 1s wrapped after being bitten. An intriguing but anecdotal observation (Walker, 1983) notes that a spider purported to be an A7gyrodes species entered the web of a red-back spider (Latrodectus hasselti Thorell) and squirted a milky substance (source unknown) onto the silk. The red-back readily ate the substance and became immobilized soon after. Such an indirect prey capture technique would provide an effective predation strategy in a small predator/larger prey system. Alternative hypotheses include stealth, the prey remaining unaware of the slowly approaching predator; or male behavioural mimicry, the predator imitating some aspect of the courtship repertoire of the prey’s male. The level and impact of A. incursus predation upon Ach. mundula populations are not clear. Data for a small sample of 17 Ach. mundula webs from Royal National Park near Sydney showed that only 2 webs harboured A. incursus individuals. By contrast 13 webs contained other presumptively kleptoparasitic/commensal A7gyrodes residents. However, Newcastle Ach. mundula webs showed A. incursus occupancy rates of approximately 50%. This suggests that A. zncursus predation may exert a considerable effect at least in small host populations. PROG. LINN. SOG. N.S.W., 111 (1), 1989 30 HOST-PREDATING ARGYRODES ACKNOWLEDGEMENTS We thank Dr H. W. Levi, Museum of Comparative Zoology, Harvard University, for confirming the generic diagnosis. References EXLINE, H., and Levi, H. W., 1962. — American spiders of the genus Argyrodes. Bull. Mus. Comp. Zool. 127: 75-204. Gray, M. R., 1983. — Spider predation on Achaearanea mundula L. Koch. Aust. Arachnol. 13: 6-8. KASTON, B. J., 1965. — Some little known aspects of spider behaviour. Amer. Midl. Nat. 73: 336-356. LARCHER, S. F., and WIsE, D. H., 1985. — Experimental studies of the interaction between a web-invading spider and two host species. J. Arachnol. 13: 43-59. ROBINSON, M. H., and ROBINSON, B., 1973. — Ecology and behaviour of the giant wood spider Nephila maculata (Fabr.) in New Guinca. Smithson. Contr. Zool. 149: 1-76. , and , 1976. — The ecology and behaviour of Nephila maculata: a supplement. Smithson. Contr. Zool. 218: 1-22. RypsTRA, A. L., 1981. — The effect of kleptoparasites on prey consumption and web relocation in a Peruvian population of the spider Nephila clavipes (L.) (Arancae: Araneidae). Ozkos 37: 179-182. TRAIL, D. S., 1980. — Predation by Argyrodes (Theridiidae) on solitary and communal spiders. Psyche 87: 349-355. VOLLRATH, F., 1979. — Behaviour of the kleptoparasitic spider Argyrodes elevatus (Araneac, Theridiidac). Anim. Behav. 27: 515-521. ——, 1981. — Energetic considerations of a spider parasite-spider host system. Rev. Azachnol. 3: 37-44. WALKER, P., 1983. — Note in: Aust. Arachnol. 14: 9. WHITEHOUSE, M. E. A., 1986. — The foraging behaviours of Argyrodes antipodiana (Theridiidac), a klepto- parasitic spider from New Zealand. New Zealand J. Zool. 13: 151-168. Wise, D. H., 1982. — Predation by a commensal spider, Argyrodes trigonum, upon its host: an experimental study. J. Arachnol. 10: 111-116. PROC. LINN. SOC. N.S.W., 111 (1), 1989 Two new Species of Amphientomidae (Insecta: Psocoptera), the first Record of the Family for Australia C. N. SMITHERS SMITHERS, C. N. Two new species of Amphientomidae (Insecta: Psocoptera), the first record of the family for Australia. Proc. Linn. Soc. N.S.W. 111 (1), 1989: 31-35. Two new species of Amphientomidae, representing the first species of the family to be recorded from Australia, are described. C. N. Smithers, Research Associate, Australian Museum, College St., Sydney, Australia 2000; manuscript received 22 June 1988, accepted for publication 24 August 1988. INTRODUCTION The psocopteran family Amphientomidae has not been recorded from Australia although Mockford (in lit.) has mentioned that the family occurs here. On this authority the family was noted as being Australian in the manuscript of the forthcoming revised edition of the ‘Insects of Australia. While sorting material in the collections of the Australian Museum two specimens have been found, each belonging to an undescribed species. They are described here. The Amphientomidae now include nearly 80 species, grouped into 18 genera. In the main they are infrequently encountered insects of warm climates. Three genera are known only from amber and there are a few species in the genus Amphientomum Pictet which are also known from amber. Current generic definitions are based mainly on venational features but these are gradually being supplemented by other characters as additional material becomes available. The Amphientomidae are unusual in that they are one of only two families in the order in which the body is clothed with scales, the other family being the unrelated Lepidopsocidae. The members of both bear a superficial resemblance to micro- lepidopterans and can easily be mistaken for them in the field. Definitions and illustrations of the main distinguishing features of the family and genera have been given in Smithers(1972); they are not repeated here. The two new species fall clearly into the genera Hemiseopsis Enderlein and Seopsts Enderlein respectively. SYSTEMATIC DESCRIPTIONS Hemuseopsis alettae sp. nov. FEMALE Coloration (in alcohol). Head and body pale testaceous, the head a little darker than body and legs. Eyes reddish brown. Morphology. Length of body: 3.1mm. Median epicranial suture fine but very dis- tinct, anterior arms absent. Pilosity of head very fine. No hairs between ommatidia. Length of flagellar segments: f,: 0.14mm; f: 0.19mm. First flagellar segment broader in basal half than in distal half, attachment to pedicel very narrow. Second flagellar seg- ment longer than first. Antennal striations very fine and dense. Scape and pedicel broad. Eyes large, not protruding from head capsule except behind. Dorsal margin con- tinuous with vertex. Seen from above eyes curve towards each other anteriorly continu- ing curvature of head. IO/D: 1.6; PO: 0.5. Lateral ocelli small, situated about a third of PROC. LINN. SOC. N.S.W., 111 (1), 1989 ae NEW SPECIES OF AMPHIENTOMIDAE (INSECTA PSOCOPTERA) distance from eye to median epicranial suture, opposite aboyt middle of eye. Lacinia (Fig. 2) curved. Maxillary palp densely beset with microtrichia. Second segment with small sensory cone and broader than third and fourth segments. Measurements of hind leg: F: 0.6mm; T: 1.0mm: t;: 0.68mm: ty: 0.11mm; ts: 0.13mm; rt: 6.2:1:1.2; ct:27,0,0. Ctenidiobothria very strong with well developed basal combs. Femur of anterior legs without cones along internal margin. Tibia of second pair of legs with external spine about half way along length in addition to usual apical spines. Femur of third leg with strong, curved, dorsal apical spine. Tibia of third leg with seven ventral spines in distal half in addition to usual apical spines. Claws with one preapical tooth and row of setulae basad of tooth. Fore wing length: 2.5mm; width: 0.78mm. Fore wing (Fig. 1) almost parallel-sided, rounded apically. Basal section of Sc ends in somewhat thickened costa. Distal section of Sc absent. R, parallel with costal margin basad of the elongate stigma- pophysis beyond which it bends to meet costal margin. Rs divides between origins of My and M3. Cu, divides opposite stigmapophysis. Hind wing with R, present. M simple. Epiproct lightly sclerotized, simply rounded behind with preapical row of setae basad of which setae are sparsely and irregularly arranged. Paraprocts simple, without clearly defined trichobothrial field but with a few large setae without patterned area around base. Subgenital plate simple. Gonapophyses (Fig. 3) with ventral valve greatly attenu- ated ending in sharp point, with ventral membranous flange. Dorsal valve broad in basal half, narrowing abruptly to form long tapering extension which matches and lies adjacent to that of ventral valve. External valve tapering to bluntly rounded end, with strongly developed dorsal lobe. MATERIAL EXAMINED. Holotype female, Falbrook, N. Singleton, New South Wales, 26.1.1979. A. S. Smithers. Holotype in Australian Museum. This species is named for my wife who collected the type specimen. DISCUSSION. Four species of Hemiseopsis have now been described, three from Africa and the present species from Australia. The genus may also occur in Jamaica (Turner, 1975). H1. alettae is smaller than H. fuellerborni (Enderlein) from Tanzania. Zaire, Angola and Zambia. In that species Rs divides opposite the origin of M3, Rs lies much further from R, and Cu, is much shorter than in H. alettae. The basal section of Rs is very much shorter than Ro,3. The genitalia of H. fuellerborni have not been described. H. machadoi Badonnel, from Angola, is only a little larger than H. alettae but can be distinguished by features of the wing venation and the female genitalia. In H. machadoi R, does not reach the wing margin and the crossvein from R, meets M basad of its separation from Rs. Ms arises opposite the division of Rs and Cu), is much shorter than in H. alettae. The ventral and dorsal valves of the gonapophyses are not strongly attenuated, reaching only as far as the end of the external valve. In H. alettae they extend far beyond the end of the external valve. In H. machadoi the lacinia is straight with a different arrangement of apical teeth from that in H. alettae. In H. obscurus Broadhead and Richards, from Kenya, several of the veins of the fore wing in the basal half are weakly differentiated. The ventral and dorsal valves of the gonapophyses are not attenuated but are blunt; the dorsal lobe of the external valve is longer than the valve itself, a condition not found in any of the species of which the genitalia have been described. The head of H. obscurus has a distinctive colour pattern not found in any of the other species. Seopsis incisa sp. nov. MALE Coloration (in alcohol). Head pale brown with brown markings. A narrow brown PROC. LINN. SOC. N.S.W., 111 (1), 1989 C. N. SMITHERS 33 “Imm Figs 1-9. Hemiseopsis alettae sp. nov. 1, female fore wing. 2, female lacinia. 3, female gonapophyses. Seopsts incisa sp. nov. 4, male femoral spines. 5, male lacinia. 6, male phallosome. 7, male fore wing. 8, male hind wing. 9, male paraproct. PROC. LINN. SOC. N.S.W., 111 (1), 1989 34 NEW SPECIES OF AMPHIENTOMIDAE (INSECTA PSOCOPTERA) mark on each epicranial plate along the ridge of the vertex; a diffuse, broad brown band across the front of the head from eye to eye; a narrow band in position usually occupied by anterior arms of epicranial suture, this broadened laterally to form a small patch just anterior to the lateral ocellus on each side; posterior half of postclypeus with a reticulate pattern in brown, part of which consists of narrow, longitudinal, parallel lines, the pattern extending further anteriorly in the midline than laterally: postclypeus pale in anterior half. Labrum pale. Lower half of genae brown. Antennae dark brown. Eyes black. Ocelli circled with dark brown. Thorax pale brown, darker along some sutures. Abdomen pale, darker dorsally on basal quarter and with suggestion of irregular seg- mental marks, laterally dark. Femora pale. Fore and middle tibiae pale in basal half, brown in distal half. Hind tibiae pale. First tarsal segment pale in basal half, darker in distal half but with pale tip. Second and third segments brown. Fore wing membrane hyaline, faintly tinged with brown, paler in distal third. Hind wing hyaline. Morphology. Length of body: 2.3mm. Median epicranial suture fine but distinct. Anterior arms indistinct but position marked by brown band. Vertex slightly curved. Length of flagellar segments: f;:0.31m; f):0.34mm. Antennal flagellum very fine, with very long setae from 7 to 15 times as long as flagellar width. Transverse striations of antennae very numerous. Eyes large, but hardly protruding. Upper margin almost level with vertex, extending a little behind head laterally. Fine, short setae between facets. Three ocelli. Lateral ocelli close to antero-medial angle of eyes, median ocellus in middle of frons so that the three ocelli are almost in a straight line. Lacinia (Fig. 5) almost straight, deeply divided at apex. Measurements of hind leg: F: 0.5mm, T: 0.98mm; t,: 0.54mm; to: 0.l1mm; t3: 0.1mm; rt: 5.4:1:1; ct: 19,0,0. Front femur with long row of short spines (Fig. 4). Claws with one preapical tooth. Fore wing length: 2.5mm; width: 0.76mm. Fore wing (Fig. 7) narrows to somewhat pointed apex. Venation as usual in genus but distal section of Sc and Cu), evanescent and less obvious than other veins. Hind wing length: 2.0mm; width: 0.6mm. Hind wing (Fig. 8) with R,; not reach- ing wing margin. IA curves distally towards hind margin. Epiproct simple, rounded behind, setose. Paraproct (Fig. 9) with finely and densely rugose, slightly raised area from which large setae arise. Bases of setae not surrounded by ‘rosette’ pattern often en- countered in barklice. Hypandrium well sclerotized, simple, posteriorly and laterally slightly upturned. Eighth sternite more heavily sclerotized than more anterior sternites and forming an anterior extension to the hypandrium. Phallosome (Fig. 6). MATERIAL EXAMINED. Holotype male, Malaise trap, AOS 125°50’E, mining camp, Mitchell Plateau, Western Australia, 9-19.v.1983. I. D. Naumann and J. C. Cardale. Holotype in Australian National Insect Collection, Canberra. DISCUSSION. Seopsis is now a genus of ten species. There are three from Sri Lanka, two from the Philippines, two from Angola and one from each of Nepal, Singapore and Aus- tralia. S. brunnea New, from Singapore, differs from S. incisa in lacking the dark post- clypeal pattern, in having a lacinia without a deeply divided apex and in lacking the row of spines on the fore femur. The male phallosome is similar to that of S. incisa in general form but differs in proportions. S. /uzonica Banks, from the Philippines, differs in having a brown head with a pale transverse band at the level of the ocelli and in having two dark marks on each side on the vertex. In S. metallops Enderlein, from Sri Lanka, the head is black or dark brown with a pale spot in the middle of the front of the head and a similar spot on each half of the vertex. The lateral ocelli are much further from the eyes than in S. incisa. The membrane of the fore wing is dark. The basal segment of the hind tarsus is more than seven times the length of the second segment, whereas it is only 5.4 times as long in S. zncisa. S. nepalensis New, from Nepal, has the posterior part of the postclypeus PROC. LINN. SOC. N.S.W., 111 (1), 1989 C.N. SMITHERS 35 pale, not dark as in S. zncisa. The fore wing 1s acuminate and there are accessory sclerites internally at the posterior ends of the male parameres. The apex of the lacinia is not deeply divided. S. pavonia Badonnel, from Angola, has the fore wing strongly acuminate and the lacinia is not deeply divided. S. termitophila Badonnel, also from Angola, has much shorter wings than S. znczsa (1.7mm as opposed to 2.5mm). The apex of the lacinia is not deeply divided and the male phallosome has a pair of pitted plates, internally, adjacent to the ends of the parameres. S. superba Hagen, from Sri Lanka, has a brown head with a darker longitudinal band on the vertex. The ocelli are close togther. The basal tarsal segment of the hind legs is very much longer than in S. incisa, being about seven times as long as the second. S. trecolor Banks, from the Philippines, has a brown head and the lateral ocelli are set well away from the eyes. It is larger than S. incisa (fore wing length 3.5mm as opposed to 2.5mm). The illustration of the wing which accom- panies the description shows no distal section to Sc, an unusual condition for this genus. S. vasantasena Enderlein, from Sri Lanka, has a brownish yellow head with four narrow, longitudinal streaks from the vertex down onto the front of the head; the postclypeus has a double, broad brown longitudinal band. The fore wing length is only 2.0mm. References SMITHERS, GC. N., 1972. — The classification and phylogeny of the Psocoptera. Aust. Mus. Mem. 14: 1-349. TURNER, B. D., 1975. — The Psocoptera of Jamaica. Trans Roy. ent. Soc. Lond. 126(4): 533-609. PROC. LINN. SOC. N.S.W., 111 (1), 1989 AG au i be 2 niger, Wai) Ao rg Hie? yy ss i i Jawa Susy 7. if he r Pi th ae Gow of a teen; wn yen et ied BAA ee Abe ve ei ivf a es ? iA a Ftd : = 7 if A Bui ee se ha 8 area sah ' nied hare i ale, Pagani eh | as ais { 19.204 alt Live eT * a ie me See’. Ure east id mua ur bid ys ins : ; Apia ig toy che peta SO A nr é 4 zs in ; Sy ab yd . a) ae ‘ pn ad? th “ wey! yas rales L wan i; Vase es v4 1 ouine, ine i oui $4 Lente Redistribution of Amino Acids and Amides during Seedling Development in Acacia iteaphylla F. Muell. (Fabaceae: Mimosoideae) ANITA B. BRADKE and DAVID R. MURRAY BRADKE, ANITA B., & MuRRAY, Davib R. Redistribution of amino acids and amides during seedling development in Acacia iteaphylla F. Muell. (Fabaccae: Mimosoideac). Proc. Linn. Soc. N.S.W. 111 (1), 1989: 37-42. The distribution of free amino compounds in the cotyledons and the organs derived from the embryonic axis has been studied in developing seedlings of Acacia iteaphylla. The results indicate that asparagine and pipecolic acid are the major forms of nitrogen translocated initially from the cotyledons to the hypocotyl and primary root. Selective transport of several ‘non-protcin’ amino acids was indicated. S-carboxycthyl- cysteine became prominent in the hypocotyl after the pipecolic acid content of the hypocotyl had declined; it appeared also in the primary leaf, but was not strongly represented in the roots. S-carboxyisopropylcysteine appeared late in the hypocotyl, and was not detected in any other part of the axis. Albizziine was not transported out of the cotyledons, but like arginine was metabolized zn situ. Anita B. Bradke and David R. Murray, Biology Department, University of Wollongong, Wollon- gong, Australia 2500 (private address D.R.M., 7 Acacia Avenue, Gwynneville, Australia 2500); manuscript received 14 June 1988, accepted for publication 24 August 1988. INTRODUCTION The cotyledons of Acacia seedlings undergo a transition from storage organs to photosynthetic organs (Ashcroft and Murray, 1979; Murray, 1981; Smith, 1981). During this transition, reserve materials appear to be mobilized in two stages. The first encom- passes the initial growth of the radicle and hypocotyl, resulting in the elevation of the cotyledons. In Acacia iteaphylla F. Muell., the dry matter content of the cotyledons is depleted by about 70% during this stage, but only a slight net decline in the protein con- tent of the cotyledons is observed (Ashcroft and Murray, 1979). In the second phase, more rapid net breakdown of protein in the cotyledons 1s associated with the synthesis of chloroplasts, as the cotyledons and then the primary leaf become effective photo- synthetic organs. It has long been known that Acacia seeds contain a high proportion of total seed nitrogen as free amino acids (Petrie, 1908, 1911). Many ‘non-protein’ amino acids have now been identified (Seneviratne and Fowden, 1968), and their distribution among species has indicated that there are four main sub-groups of Acacia throughout the world (Evans e¢ al., 1977; Murray, 1986b). In view of the early slow rate of net decline in the protein content of cotyledons, Ashcroft and Murray (1979) suggested that non-protein amino acids might represent the earliest mobilized form of nitrogenous reserve. Evidence has now been obtained that some, but not all, of the non-protein amino acids are transmitted to the axis, together with newly-synthesized asparagine. MATERIALS AND METHODS The seeds of Acacia iteaphylla were from the same batch studied previously (Ashcroft and Murray, 1979). To permit uniform imbibition, the seedcoats were cut with a razor blade at the end furthest from the embryonic axis. Except for seeds imbibed for 24h, which were placed in Petri dishes between moist Whatman No. 1 filter paper at 23°C (Krishna and Murray, 1988), seeds were placed in trays of sandy soil in a glass house, PROC. LINN. SOC. N.S.W., 111 (1), 1989 38 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS with day temperatures around 30°C. At intervals up to 14 days, groups of 10 to 20 uniform seedlings were removed, dismembered and analysed. Extracts from cotyledons, radicles and hypocotyls were prepared using a chilled mortar and pestle, acid-washed sand, and medium consisting of 235mM K* phosphate (pH 7.5), 0.5mM 2-mercaptoethanol and 0.03% (w/v) Triton X-100 (Murray and Kennedy, 1980; Murray, 1983). The ratio of medium to tissue was 5:1 (mL per g fresh weight). The homogenates were centrifuged for 4min at 9,000g in a Microfuge and the supernatants removed. These were treated with ethanol (4:1, v/v) and insoluble material removed by centrifugation (Murray and McGee, 1986). Smaller samples (whole axes; balance of shoot) were extracted directly with ethanol (4:1, v/v). Aliquots of the ethanol- soluble fractions were assayed for amino nitrogen content by the procedure of Schnar- renberger et al. (1972), using L-serine as a reference standard (Murray, 1983). Ethanol-soluble fractions were dried in a rotary evaporator, then redissolved in 0.5 to 1.0mL of 70% (v/v) ethanol, and applied to the origins of Whatman No. | papers pre- pared for 2-dimensional descending chromatography (Murray, 1983). Extracts avail- able only in small quantities were applied without prior concentration. The solvents employed were 80% (w/v) phenol-water plus ammonia (200:1, v/v) in the first dimension, then either n-butanol: acetic acid: water 12:3:5 (v/v) or n-butanol: propionic acid: water 6:3:4 (v/v) in the second (Murray e¢ al., 1971; Murray, 1983). At least two chromatograms were run for each extract. Amino compounds were detected by their reaction with ninhydrin, and identified by comparison of their positions with those of authentic compounds. Information on the chromatographic behaviour of non-protein amino acids was kindly provided by Dr C. S. Evans and Prof. E. A. Bell. Authentic albizziine was purchased from Aldrich; other amino acids and amides were from Sigma Chemical Co. RESULTS The rate of seedling growth was faster than in the previous study because of higher temperatures and longer day-length. The 4-day, 8-day and 14-day stages chosen for analysis closely resembled the 8-day, 15-day and > 21-day stages described previously (Ashcroft and Murray, 1979; Murray, 1981). At the 1-day stage, the amino nitrogen con- tent of the cotyledons effectively represents that of the whole embryo (Fig. 1), as the axis was too small for accurate analysis (less than Img fresh weight). The amount of free amino nitrogen in the whole seedling progressively increased, doubling by 14 days (Fig. 1). Within the cotyledons, the amino nitrogen content did not alter substantially until between 8 and 14 days, when it declined. Within the axis, the hypocotyl gained a much higher proportion of the amino nitrogen exported by the cotyledons than did the radicle. The amino acids and amides present in Acacia seedling tissues were identified as shown in Fig. 2. Changes in their relative abundance in the cotyledons are shown in Table 1. Many of the major nitrogenous solutes stored in the cotyledons initially are still among the most prominent forms present after 14 days, when the total free amino nitro- gen content of the cotyledons had declined by 40% (Fig. 1). The content of asparagine increased, while the contents of albizziine, glycine, arginine, pipecolic acid, and finally glutamate and alanine, declined (Table 1). The distribution of amino compounds in the roots and hypocotyls of seedlings aged 8 and 14 days is shown in Table 2. Pipecolic acid and asparagine were the most abundant forms of free amino nitrogen in both the roots and the hypocotyl] of the 8-day-old seed- lings. At this stage, S-carboxyethylcysteine was prominent in the hypocotyl, but was not detectable in the roots. At the 14-day stage, both S-carboxyethylcysteine and S-carboxy- isopropylcysteine became prominent in the hypocotyl. The content of asparagine was PROC. LINN. SOC. N.S.W., 111 (1), 1989 ANITA B. BRADKE AND DAVID R. MURRAY 39 Mmoles amino N oOo + NY OO bh HO See A R DAYS: 1 4 8 14 Fig. 1. Changes in the distribution of free amino nitrogen in the developing seedling of Acacia iteaphylla. C = cotyledon pair; A = whole embryonic axis; R = radicle or root system; H = hypocotyl; S = shoot system above the cotyledons. NOISNIWIG puZg PIPECOLIC ACID aS = ana (ae ORIGIN 1st DIMENSION Fig. 2. The positions and identities of amino compounds separated from cotyledons of Acacia iteaphylla by 2- dimensional descending paper chromatography. Solvent 1, phenol-water-ammonia; solvent 2, n-butanol- acetic acid-water (sec Materials and Mcthods). Abbreviations are standard, plus S-CEC = S- carboxyethylcysteine; S-CIC = S-carboxyisopropylcysteine. ; PROC. LINN. SOC. N.S.W,, 111 (1), 1989 40 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS TABLE 1 Changes in the Distribution of Free Amino Acids and Amides in the Cotyledons of Acacia iteaphylla Seedlings Age of scedling (days) Compound 3 4 8 14 S-carboxycthylcysteine ++++ ++++ ++++ +++ S-carboxyisopropyleysteine +++ ++44 ++4+4+ 45 te fe Pipecolic acid +++ ++4++ +44 de Le Albizziine + asparagine pal ++ Rae soup a2 Glutamine ++ ++ See eet A tt Aspartate tr ++ tr fe Glutamate +++ +++ fe fe fe Serine ++ + + ++ 4 ae Glycine ++ + + + Alanine ++ +++ ++ tr Valine + tr tr tr Leucine + Isoleucine + tr tr tr Lysine tr tr tr tr Arginine ++ + tr n.d. tr, trace; n.d., not detected. albizziine predominant; asparagine predominant — sec text. TABLE 2 Changes in the Distribution of Free Amino Acids and Amides in the Roots and Hypocotyl of Acacia iteaphylla Seedlings Roots Hypocotyl Compound 8-day 14 day 8-day 14-day S-carboxycthylcysteine n.d. + ++ +++ S-carboxyisopropylcysteine n.d. n.d. n.d. ++ Pipecolic acid +++ ++ ++4++4+ + Asparagine +++ ++ ++4++4+ +++ Glutamine ++ n.d. ++ n.d. Aspartate + + ++ + Glutamate + n.d. fp ae tr Serine + ++ ++ + Glycine tr ++ + n.d. Alanine ++ + + +++ tr Valine ++ n.d. + n.d. Leucine + Isoleucine tr n.d. + n.d. Threonine n.d. n.d. tr n.d. tr, trace; n.d., not detected. maintained, but the contents of pipecolic acid and alanine declined sharply (Table 2). In the roots of the 14-day-old seedlings, some S-carboxyethylcysteine was present, but S- carboxyisopropylcysteine was not detected. In the balance of the shoots from these seedlings (mainly the primary leaf at 14 days), serine and S-carboxyethylcysteine were most prominent. Pipecolic acid was also detected, along with aspartate, glycine, alanine, glutamate and glutamine, but asparagine, albizziine and S-carboxyisopropylcysteine were not detected. PROC. LINN. SOC. N.S.W.,, 111 (1), 1989 ANITA B. BRADKE AND DAVID R. MURRAY 41 DISCUSSION The net increase in free amino nitrogen in the seedling between 1 and 8 days was 2.55umol (Fig. 1), which is within the maximum amount that could be met from limited breakdown of proteins in the cotyledons (Ashcroft and Murray, 1979). Our observations are consistent with the transport of selected forms of free amino nitrogen from the cotyledons to the hypocotyl, the root, and later the shoot, with the content of free amino nitrogen maintained close to 2umol per cotyledon by proteolysis until after 8 days (Fig. 1). It must be noted that it is not possible for any uptake of exogenous nitrogen sources to have occurred through the roots; the axis-derived organs are gaining nitrogen entirely at the expense of the cotyledons. Since the only way that the cotyledons can export nitrogenous solutes is through the phloem (Guardiola and Sutcliffe, 1972; Murray, 1984), an important question raised is whether the different organs of the axis are selective in the nitrogenous solutes they import, and if so, whether restrictions apply to phloem loading or unloading of in- dividual compounds. In seedlings of jack bean (Canavalia ensiformis |L.] DC), Rosenthal and Rhodes (1984) have demonstrated that the non-protein amino acid L-canavanine is transported only to the above-ground parts of the shoot — none is translocated to the radicle. Of all the non-protein amino acids potentially available from the cotyledons, only pipecolic acid appeared to be imported by very young roots of Acacza iteaphylla seedlings. Pipecolic acid and asparagine were abundant early in both the roots and the hypocoty]; quantitatively, they are the most important forms of nitrogen initially transferred from the cotyledons to the axis (Table 2). Outside the cotyledons, S-carboxyisopropylcysteine was detected only in the hypocotyl, relatively late in development (Table 2), whereas transport of the other derivative of cysteine was less restricted. S-carboxyethylcysteine became one of the more prominent nitrogenous solutes of the hypocotyl, and appeared also in the roots (late) and the primary leaf. The serine present in the primary leaf has probably accumulated from at least two biosynthetic pathways operating in the leaf itself (Murray, 1986a). Arginine was not transported from the cotyledons (Table 2), but converted to other nitrogenous solutes in situ (Table 1). Urease was detected immunochemically in cotyledon extracts (for methods see Murray and Knox, 1977), which is consistent with this interpretation. Finally, what is the role of albizziine (L-a-amino-$-ureidopropionic acid), a com- pound whose synthesis in Acacia cotyledons is known to be confined to seed development (Seneviratne and Fowden, 1968)? In our chromatographic systems, albizziine and asparagine could not be resolved (Fig. 2). This is in agreement with data on the mobility of albizziine reported by Cooper and Meister (1973). However, the colour of the reaction product with ninhydrin produced by authentic albizziine after paper chromatography was always mauve, whereas that produced by asparagine was always brown. On chromatograms where authentic albizziine and asparagine were loaded together, the final colour reflected the abundance of each compound; a brown colour consistently indicated the predominance of asparagine. Initially albizziine is the prominent component of the combined albizziine- asparagine area from cotyledon extracts, but with increasing time following germi- nation, asparagine predominates. When detected on chromatograms of Acacia root and hypocotyl extracts, the joint region was always brown, and if the unstained area was eluted from replicate chromatograms and hydrolysed with HCl, a ninhydrin positive product with the same mobility as aspartate was recovered. We have therefore con- cluded that asparagine is a major nitrogenous solute translocated to the roots and PROC. LINN. SOC. N.S.W., 111 (1), 1989 42 MOBILIZATION OF NITROGEN IN ACACIA SEEDLINGS hypocotyl (Table 2). This is in agreement with a transport function for newly synthe- sized asparagine determined for cotyledons of light-grown pea (Melcher, 1983) and peanut seedlings (Peoples et al., 1986). In Acacia cotyledons albizziine, like arginine, is converted to translocated forms of nitrogen, predominantly asparagine. It remains to be determined exactly how albizziine is metabolized, and whether, while present in cotyledons, it has any deterrent function against larvae of Australian insects. ACKNOWLEDGEMENTS The financial support of the Joyce W. Vickery Scientific Research Fund (Linnean Society of N.S.W.) is gratefully acknowledged. We also thank Dr C. S. Evans and Prof. E. A. Bell for helpful information and advice. References ASHCROEFT, W. J., and Murray, D. R., 1979. — The dual functions of the cotyledons of Acacia iteaphylla F. Mucll. (Mimosoideac). Aust. J. Bot. 27: 343-352. Cooper, A. J. L., and MEISTER, A., 1973. — Action of liver glutamine transaminase and L-amino acid oxidase on several glutamine analogs. J. Biol. Chem. 248: 8499-8505. EVANS, C. S., QURESHI, M. Y., and BELL, E. A., 1977. — Free amino acids in the seeds of Acacia spccics. Phytochem. 16: 565-570. GUARDIOLA, J. L., and SUTCLIFFE, J. F., 1972. — Transport of materials from the cotyledons during germi- nation of seeds of the garden pea (Pisum sativum L.). J. Exp. Bot. 23: 322-337. KRISHNA, T. G., and MuRRAY, D. R., 1988. — Effects of cycloheximide and actinomycin D on glycosidase activities in the cotyledons of legume seeds following imbibition. J. Plant Physiol. 132: 745-749. MELCHER, I. M., 1983. — Changes in nitrogen-containing compounds of the garden pea (Pisum sativum L.) during germination. Z. Pflanzenphysiol. 112: 95-102. Murray, D. R., 1981. — Functions of the cotyledons in Acacia. Aust. Plants 11: 65-66. ——.,, 1983. — Changes in free amino acid and amide composition during fruit and seed development of garden pea, Pisum sativum L. New Phytol. 93: 33-41. ——, 1984. — Axis-cotyledon relationships during reserve mobilization. Jn MurRRay, D. R., (ed.), Seed Physiology. Volume 2. Germination and Reserve Mobilization: 247-280. Sydney: Academic Press. —, 1986a. — Amino acid and amide metabolism in the hulls and seeds of developing fruits of garden pea, Pisum sativum L. 1V. Alanine. New Phytol. 104: 395-406. ——.,, 1986b. — Sced dispersal by water. Jn MuRRAY, D. R., (ed.), Seed Dispersal: 49-85. Sydney: Academic Press. , and KENNEDY, I. R., 1980. — Changes in activitics of enzymes of nitrogen metabolism in sccdcoats and cotyledons during embryo development in pea seeds. Plant Physiol. 66: 782-786. , and Knox, R. B., 1977. — Immunofluorescent localization of urease in the cotyledons of jack bean, Canavalia ensiformis. J. Cell Sci. 26: 9-18. —-, and McGrr, C. M., 1986. — Sced protein content of Australian species of Acacia. Proc. Linn. Soc. N.S.W. 108: 187-190. ——., GIOVANELLI, J., and SMILLIE, R. M., 1971. — Photometabolism of glycolate by Euglena gracilis. Aust. J. Bool. Sci. 24; 23-33. PEOPLES, M. B., PATE, J. S., ATKINS, C. A., and BERGERSEN, F. J., 1986. — Nitrogen nutrition and xylem sap composition of peanut (Arachis hypogaea L. cv. Virginia Bunch). Plant Physiol. 82: 946-951. PETRIE, J. M., 1908. — The role of nitrogen and its compounds in plant metabolism. Part 11. The non-protein nitrogen in seeds. Proc. Linn. Soc. N.S.W. 33: 835-844. ——,, 1911. — The role of nitrogen in plant metabolism. Part 111. The distribution of nitrogen in Acacza seeds. Proc. Linn. Soc. N.S.W. 36: 97-126. ROSENTHAL, G. A., and RHODES, D., 1984. — L-Canavanine transport and utilization in developing jack bean, Canavalia ensiformis (L.) DC [Leguminosae]. Plant Physiol. 76: 541-544. SCHNARRENBERGER, C., OESER, A., and TOLBERT, N. E., 1972. — Isolation of protein bodies on sucrose gradients. Planta 104: 185-194. SENEVIRATNE, A. S., and FOWDEN, L., 1968. — Diamino acid metabolism in plants with special reference to a,@-diaminopropionic acid. Phytochem. 7: 1047-1056. SMITH, D. L., 1981. — Cotyledons of the Leguminosae. Jn POLHILL, R. M., and RAVEN, P. H., (eds), Advances in Legume Systematics. Part 2: 927-940. Kew: Royal Botanic Gardens. PROC. LINN. SOC. N.S.W., 111 (1), 1989 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 111 NUMBER 2 John Vaughan Thompson, F.L.S. HUGH CAMPBELL (Communicated by T. G. VALLANCE) CAMPBELL, HUGH. John Vaughan Thompson, F.L.S. Proc. Linn. Soc. N.S.W. 111 (2), 1989: 45-64. John Vaughan Thompson (1779-1847) was a surgeon in the British Army Medical Service who made his name as a ‘gentleman naturalist. He became a Fellow of the Linnean Society in 1810 in recognition of his work in botany, but his great contribution to science was his discovery of metamorphosis in the Crustacea: a discovery which was received with disbelief and hostility in the 1830s, and not fully accepted until after Thompson died. He spent the last eleven years of his life in New South Wales, eight of them in charge of Convict and Military Medical Services. He failed as an adminis- trator, and most Australian historians have not only blamed him unduly for his failure but have also neglected the scientific achievements which more than compensate for it. His life is an interesting study of the difficulties faced by an amateur scientist working alone without the support of personal wealth or powerful friends. N. H. Campbell, 19 Brushy Creek Road, Lenah Valley, Australia 7008; manuscript received 19 July 1988, accepted for publication 23 November 1988. INTRODUCTION — GIVING J. V., THOMPSON HIs DUE John Vaughan Thompson (1779-1847) distinguished himself as a zoologist and botanist while quietly earning his living as an army surgeon in England, the West Indies, Mauritius, Madagascar, Ireland and Australia. He has been treated generously by his recent English biographers, who refer to his achievements before he went to Aus- tralia. Australians have been less kind to him: if they have noticed him at all, it is only in passing. For instance, although he spent his last eleven years in New South Wales, eight of them in charge of the convict and military medical services in the colony, the ADB contains no entry about him in his own right: he is mentioned only as a sort of béte nozre in other men’s lives. It is true that he failed as a colonial administrator, and only natural that local historians should have reported accordingly. But most did not inquire further, and so overlooked both his zealous service during forty-five years as an army surgeon and his wide-ranging work in natural history, especially his discoveries about marine in- vertebrates. What is more, the writer believes they made their unflattering picture even worse by blaming him too much for his difficulties in New South Wales. As a result, biographers sympathetic to Thompson have usually thought it best to dismiss his years there in a single sentence and concentrate on the rest of his life. This paper is an attempt to repair the large gaps and unfair biases in what has hitherto been written about him. There are other reasons, too, for the discrepancies. First, Thompson was inconsis- tent in himself: illness and bitterness changed him during the course of his service in New South Wales. Second, earlier biographers do not mention the Colonial Office files, reported for the first time in this paper, which contain a number of details of his personal life. Hence, they do not explain some of his actions. Third, several of his discoveries in natural history soon became such common knowledge that his later biographers forgot his part in discovering them. Wheeler (1970) pointed out that most of Thompson's important achievements were recognized too late for him to enjoy the praise: he had that sort of luck all his life. By his own account, fortune first failed him in 1778, the year before he was born, when his father lost all his property in North America. In one part of a ‘memorial’ seeking a grant PROC. LINN. SOC. N.S.W., 111 (2), 1989 46 JOHN VAUGHAN THOMPSON, F.L:S. of land in New South Wales, Thompson (1835) explained the setback: ‘Memorialist’s father . . . lost to his family by attachment to the Loyalist cause and the unfortunate result of the American War, a grant of land to the amount of 4,000 acres situated on the Mohawk River beyond Albany, . . . together with a good deal of other property, houses, &c at New York and Long Island, and a considerable stock of firing provided for the army under Sir John Vaughan, for which he was contractor — the above grants being given as the reward of services as an officer of the 96th Regiment at the taking of Pt Havanna [sic] &c was totally lost by the precipitation with which he was obliged to fly when the King’s troops took their final departure from Sandy Hook’ .* It was typical of Thompson to expect the government to compensate him for the loss his Loyalist father had suffered in the American War of Independence more than fifty years earlier — a loss he had probably heard of many times as a boy. But, as we shall see, to the end of his life, he remained surprisingly naive in political matters. On 19 November 1779, a year after his father fled from Sandy Hook, John Vaughan Thompson was born at Berwick-on-Iweed. His mother was Jane Hall (International Genealogical Index, 1981). Thompson grew up there, and gave that address as late as 1817. But he left home much earlier: the son of a dispossessed Loyalist army officer had to make his own way in the world. He studied medicine at the University of Edinburgh in the sessions of 1797-8 and 1798-9, enrolling in anatomy, surgery, obstetrics, chemistry and botany (Edinb. Univ., 1797-8); and on 3 March 1799, soon after his nineteenth birthday, he was appointed Assistant Surgeon to the Prince of Wales Fencibles. Without taking his degree, he accompanied the regiment to Gibraltar in December 1799. THE WEST INDIES, 1800-1809 Appointment as Army Surgeon On 3 July 1800 he became Assistant Surgeon of the 37th Foot, and embarked with them for the West Indies and Guiana to take part in the war against the French and Dutch: records show that he was engaged in the taking of Berbice and Demerara in Guiana. On 25 June 1803 he was promoted to Surgeon in the regiment. During his six years residence in Trinidad, with occasional visits to Grenada and St Vincent, he found time, while carrying out his military duties, to study the natural history of subjects as diverse as land crabs and ginger. There is evidence that he paid at least one visit to England during his service in the West Indies. When he asked Lord Seaforth, F.L.S. and recently retired Governor of Barbados, to present his paper about Kaempferia to the Linnean Society, he headed his covering letter ‘London, April 7th, 1807’ (Linn. Soc., 1807). He took advantage of being in England that year to engage in other literary activity: he published his ‘Catalogue of Plants in the vicinity of Berwick-on- Tweed’ (a small octavo of 132 pages) (Thompson, 1807a), and had papers on Kaempferra (Thompson, 1807b) and Piper (Thompson, 1808) read to the Linnean Society. Early Interest in Botany Britten (1912), reviewing his first publication, wrote: “Thompson must early have acquired considerable proficiency in botany. His Catalogue of Plants ... Berwick upon Tweed was prepared before he left England — i.e. at or before the age of twenty — and * The Encyclopedia Americana (1983) explains the reference to the capture of Havana in 1762; the DNB (1900a) entry on Sir John Vaughan gives an account of his service on the British side in the American War of Indepen- dence and his appointment as Governor of Berwick-on-Iweed, where Thompson’s father appears to have joined him, a possible connnection by a marriage in 1765 being suggested by the International Genealogical Index (1981); and the EB gives an account of the British flight from Sandy Hook in 1778. PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 47 shows a very complete knowledge of the plants of that region and of the literature of the period: the pretty coloured group which adorns the title page and a plate (also coloured) and dissections show him to have been a capable artist’. In the same article, Britten points out that a paper on British birds has been wrongly attributed to Thompson (DNB, 1900b) because of a false entry in Royal Society (1871). Thompson left the West Indies for good and returned to England in 1809. A year earlier he had been made an Associate of the Linnean Society, and in November 1809, four members signed the following certificate: ‘John Vaughan Thompson, Esq., Surgeon of his Majesties [sic] 37th Regiment of Foot, & A.L.S., a Gentleman well versed in the study of Botany, being desirous of becoming a Fellow of the Linnean Society of London, we, the undersigned do from our personal knowledge beg leave to recommend him as likely to become worthy of that honour (Linn. Soc., 1809).* He was elected F.L.S. on 6 February 1810. Stebbing (1910), writing on the centenary of the occasion, described his election as ‘prophetic insight’, recognizing that Thompson's most important work was still to come. On 3 March 1812, two of his papers were read to the Society — one about an unusual Pouched Rat he had observed in Trinidad (Thompson, 1813) and the other, reported by Wheeler (1970), on his observations of the genus Myrti there. SERVICE IN MAURITIUS AND MADAGASCAR, 1812-1816 Shortly afterwards, he was posted to Mauritius and Madagascar. On Christmas Day, 1812, he was appointed staff surgeon, though his official duties in the Mascarene Islands are not now known. It has been said that in two consecutive years he worked on ‘Introducing vaccine innoculation to Madagascar, and that for some time he was officially styled ‘Government Agent for Madagascar’. In his 1835 letter seeking a land grant (Thompson, 1835), he himself refers to his appointment instructions and his jour- nal ‘sent to Governor Sir R. T. Farquhar and now, he presumes, in the Colonial Office’. Neither the instructions nor the journal have been found. He claims (Thompson, 1835) that: ‘amongst the important accessions to the culture of the Mauritius, [he] added 8 varieties of that estimable food, the plaintain, 10 valuable varieties of banana, 12 varieties of sugar cane, most of them vastly superior to those in common cultivation, 9 varieties of tobacco, 7 varieties of the indigo plant and 11 varieties of ordinary and upland rice’. He also writes of: ‘His efficient service as Civil Agent at Madagascar for the years 1814-15 a. During the period of his Agency he put a total stop to the traffic in slaves from those parts under his control. b. Repressed several efforts of rival chiefs to disturb the general tranquility [sic] by war. c. Introduced vaccine innoculation amongst the natives. d. Completed a survey of the extensive harbours of Loquez without loss of a single life. e. Paved the way for a friendly intercourse with the natives by a kind and generous treat- ment diametrically opposed to that of the French, our predecessors. f. Introduced and taught the cultivation of the potato and several other culinary vegetables’. Vaughan (1953) refers to a 1902 work by Grandidier in which Thompson was credited with introducing Albizzza lebbek (‘bois noir’) to the treeless plains of Madagascar, having included a sack of seeds among his presents to the island’s king, Radama I. The reference is further evidence that Thompson was engaged in some sort of diplomatic service in Madagascar. * Apparently wishing, on second thoughts, to show their wholehearted endorsement of an Associate, the sponsors struck out ‘likely to become’. PROC. LINN. SOC. N.S.W., 111 (2), 1989 48 JOHN VAUGHAN THOMPSON, F.L:S. Thompson says that eventually he was ‘attacked by the formidable remittent fever of the country which obliged him to relinquish a very lucrative post to return to Europe at great expense and finally to go on the half pay of his then rank of surgeon’. In his informative ‘land grant letter, he also makes a particular point of mentioning that he had ‘at the request of Governor Farquhar, drawn up and published at the Mauritius a systematic catalogue of all the plants cultivated in the three Government Gardens of Pamplemousses, Mon Plaisir and Reduit, which cost [him] much labour, numerous journeys and a great deal of patient investigation’. The title page of the Catalogue of Exotic Plants Introduced to Mauntius (Thompson?, 1816) does not show Thompson as the author. R. E. Vaughan (1953), however, based an article entitled A Forgotten Work of John Vaughan Thompson on yet another letter, which Thompson had written about the Catalogue in 1838. In that letter, he claimed to be the author of the work, which had in fact been published anonymously in 1816, shortly after he had left for England. He asserts that someone had suppressed the title page naming him as author, which had been ready for the press when he sailed, and replaced it with another. The 1838 letter was an appeal to the President of ‘La Société d’Histoire Naturelle de Maurice’ asking the Society to credit Thompson with the authorship. In the event, the President did nothing to advance Thompson's claim, partly because he thought that the Catalogue, based as it was on the Linnaean ‘systeme sexuel, was out of date by 1838. Regardless of authorship, ‘this was the first work on the plants of the island to be published locally, and is useful in listing the dates of introduction of many plants grown in the Botanic Gardens and elsewhere. It shows Thompson’s interest in the importation of useful plants, an interest he shared with many. other colonists of the time’ (Wheeler, 1970). In fact, it contains references to a number of plants introduced by Thompson himself. Barnwell (1941) lists among these the honeysuckle, the jonquil, Guernsey and other lilies, Madagascar arrowroot and tobacco, weeping willow, walnut, chestnut, horse-chestnut and American lime; he also records that Thompson’s name is inscribed on the Liénard Obelisk in the Pamplemousses Garden. But this first taste of having his scientific efforts and their worth doubted probably influenced Thompson’s later decision to publish his own work. Britten (1912) reports that during this period he also ‘sent dried plants to Robert Brown, which are in the National Herbarium’, later to become part of the British Museum (Natural History). Reviewing his service between 1812 and 1816, Thompson later complained of the ‘loss of property at the Madagascar and the Mauritius, being involved in the latter by the burning of Port Louis* and the extensive bankrupcies [sic] which occurred shortly after his departure for Europe on sick leave’, and of the ‘loss and expense entailed by the great length to which the regnal|[?] of the Madagascar was extended’. When Thompson left Mauritius, he was thirty-seven. In the year of his departure, he suffered physically from the onset of malaria, financially from the fire at Port Louis, and psychologically from lack of recognition of his efforts. And his hardship was com- pounded because illness forced him to go on half pay for two years from 10 June 1817 (Johnston, 1917). He had the consolation, however, of knowing that his paper on Mus anomalis, which had been read before he left England, had been published by the Linnean Society in 1813 (Thompson, 1813). The paper, accompanied by a plate which illustrates his *“A fire broke out in the commercial section of Port Louis [in 1815]. Efforts to extinguish it were unsuccessful and all the principal shops and warchouses were destroyed. Seven hundred houses were burned down, and millions of rupees of property lost. [Governor] Farquhar acted quickly and efficiently. People were clothed and fed and no one died of hunger, although many were ruined financially . . .’(Mannick, 1979). PROC. LINN. SOC. N.SW.,, 111 (2), 1989 HUGH CAMPBELL 49 meticulous drawing, contains a revealing statement of his views about scientific classifi- cation: ‘But when we examine into nature with due attention, we find she delights to mock the vain efforts of mortals to shackle and confine her within the bounds of generic characters, which are found to run so into each other as to render all attempts at method more or less imperfect. This animal must remain an anomaly in the family’. His curiosity about natural history was not dampened. He had taken advantage of his time in Mauritius to study ‘the famous extinct Mascarene birds’, although he waited until 1829 to publish his Contributions towards the natural history of the Dodo . . . (Thompson, 1829). More immediately, as Wheeler (1970) records, on the way home he observed south of Madagascar ‘a puzzling luminosity in the sea. He trailed a muslin hoop over the stern of the ship and caught a profusion of small animals hitherto invisible in the water’. Wheeler goes on to note that “Thompson has been credited with being the first to use a plankton net, and there is little doubt that his use of it in late July or August 1816 was his own idea entirely: but he was anticipated by John Cranch, who used a similar tow net on [Captain Tuckey’s voyage to] the River Zaire (or Congo) in April 1816. Back in Berwick-on-Iweed on half pay, Thompson returned to medical school. He enrolled for the 1816-17 session at the University of Edinburgh in Clinical Medicine and Materia Medica. He paid 4/6d for lectures in Materia Medica from Professor James Home; among others on his page of the fees list were two Royal Navy surgeons and another army surgeon (Edinb. Univ., 1816). ARMY MEDICAL SERVICE AND MARINE STUDIES AT CORK, 1819-1835 The next record of Thompson’s movements appears when he was restored to full pay (on exchange) on 25 May 1819 (Johnston, 1917). By that time he was in Cork, and for the ensuing eleven years he remained on full pay in posts such as district medical officer at Cork and surgeon of the Cork Recruiting District. There is evidence that in 1819 he gave a course of lectures at the Cork Institute on the science of Botany (Wheeler, 1977/5) Presumably he married soon after his return to England in 1816. The advice of his departure from Mauritius in June 1816 (Gazette de l’Isle Maurice, 1816) suggests that he had no wife then. Previous biographers have said nothing about his private life and family; Wheeler (1970), the latest, said, ‘It is not known whether he married’ But it is a subject on which Thompson’s (1835) revealing letter throws some light. In it, he describes himself as ‘having a young family of six children, four girls and two boys, whose prospects have been materially affected’ by the reverses in North America and Mauritius which he mentioned in the letter. Tantalizingly, the shipping record shows that, when he sailed for Hobart Town in 1835, he was accompanied by his wife and four children, the eldest sixteen (Colonial Office, 1836a): for some reason, two of the girls (probably older than 16) did not go. Thompson remained at Cork until 1835. During his years there he made those discoveries of the life histories of the marine invertebrates in the Cove [Cobh] for which he is chiefly remembered. He announced the principal ones in the following order during the brief period between 1827 and 1830: the discovery of Pentacrinus europaeus (Thompson, 1827); t the discovery of metamorphosis in the development of most Crustacea (Thompson, 1828); the classification and life history of barnacles (Thompson, 1830a); and the discovery of the animals he called Polyzoa (Thompson, 1830b). As the DNB (1900b) has it, ‘Our present conceptions of the structure of these forms, of PROG. LINN. SOG. N.S.W., 111 (2), 1989 50 JOHN VAUGHAN THOMPSON, F.LS. their zoological position, and of the metamorphoses which they undergo, date from Thompson's papers. His achievements resulted from his appreciation of the need for practical obser- vation, his capacity for detecting and recording detail and, above all, his ability to realize the implications of what he saw. They were all the more remarkable because he worked alone, without the support of a university or other centre for scientific develop- ment, or the collaboration, it seems, of even a single colleague. Hence, for want of people to rally round him in his lifetime, his name is not well known today, even among specialists in marine zoology. Although his discoveries about marine invertebrates received a good deal of attention when he announced them, biologists soon began to take them for granted: and if scientists subsequently wondered who made those dis- coveries, they often attributed them to later workers. Thompson announced his discoveries in a series of memoirs which he published at his own expense. Although nowhere does he say so, it is likely that his experience with his Catalogue of Exotic Plants in Mauritius made him wary of trusting his discoveries to others until he had published them. Pentacrinus europaeus The DNB (1900b), in its reference to his first discovery, the Memoir on Pentacrinus europaeus (Thompson, 1827), records that it: ‘announced the presence of a stalked crinoid in the seas of the British Isles: until then crinoids (feather stars or sea-lilies) were known only from the West Indies. Thompson also revealed that the crinoidea were really ‘radiata’; and (as shown more fully by a second paper in New Philosophical Transactions, Edinburgh 1836) that this pentacrinus was really the young stage of the antedon, or feather star’. The discovery of this animal and Thompson’s conclusions about the stages in its life history drew the attention of zoologists in France, Germany and elsewhere to his work, and many of his succeeding papers were translated or abstracted into journals abroad. Thompson announced his other principal discoveries in a series of five scarce pamphlets, now reprinted in facsimile, which he published under the title of Zoological Researches and Illustrations (Thompson, 1828-34). In the ight of Thompson’s later appointment to Sydney, it is interesting to note that a copy of the first Researches was sent to New South Wales, endorsed ‘W. S. McLeay, Esq., with the author’s compliments. ‘He was also the donor of seeds of cotton to A. McLeay in Sydney, as shown by the Seed Book’ (Fletcher, 1920). In fact, Thompson corresponded regularly — and evidently was on familiar terms — with Alexander Macleay, the Treasurer of the Linnean Society of London from 1798 until 1825, and thereafter Colonial Secretary of New South Wales (Wheeler, 1970). Macleay, of course, was one of the four who had nominated him for F.L.S. in 1809. Metamorphosis of Crustacea Thompson published the first issue of his Researches at Cork in September 1828, price 3s.6d. ‘It announced what is probably his most important contribution to zoology: the discovery that certain planktonic forms of crustacean, then known by the generic name Zoea, undergo changes of form (metamorphoses) until they become recognizable as the young of the European edible crab (Cancer pagurus) (Wheeler, 1970). The full title of the first memoir was: On the Metamorphoses of the Crustacea, and on Zoea, exposing their singular structure and demonstrating that they are not, as has been supposed, a peculiar genus, but the Larva of Crustacea!! (Thompson, 1828). Wheeler (1975) points out that ‘the two exclamation marks as well as his text showed that he appreciated the revolutionary PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 51 nature of his discovery, but it is doubtful whether he could have anticipated the furore that his important discovery was to invoke among the zoological establishment’. Wheeler (1975) goes on to explain how Thompson’s discovery challenged one of the basic tenets of contemporary systematic zoology: “The essence of the controversy lay in the distinction made by systematists between the classes /nsecta and Crustacea. The insects, as was easy to observe, went through a series of metamorphoses to emerge as an imago, or perfect insect. The crustaceans, less easy to observe, were not known to develop through metamorphosis; consequently they were assumed not to do so. This difference was one of the principal characters distinguishing the classes’ Wheeler’s summary of Thompson’s memoir is simple and elegant: “Thompson’s demolition of the character was based on practical observation. In 1816, while on the return journey from Mauritius, he had . . . captured small planktonic animals, which resembled species of the crustacean genus Zoea, described by earlier naturalists from mid-ocean. In the spring of 1822, Thompson “to his great surprise” found a considerable abundance of Zoeas in the harbour of Cove [Cobh], and in 1823 he succeeded in keeping a large specimen in fresh sea water between 14 May and 15 June, when it died in the process of casting its skin. He noted that the form of the Zoea was totally changed and that the limbs that had disengaged from the skin resembled those of the decapod crusta- ceans (crabs, lobsters, shrimps, etc.) in having four long walking legs and an anterior pair armed with pincers. The final confirmation was obtained four years later, in June 1827, when Thompson succeeded in hatching the eggs of the edible crab (Cancer pagurus) to find zoea larvae. From these observations he asserted that the decapod crustaceans generally undergo metamorphosis during their development. Not only did his announcement render invalid ong of the major distinctions between the Crustacea and the Insecta, it accounted for the anomalous Zoea specimens of earlier naturalists, and it also explained previously unaccountable phenomena such as the migration to the sea of land crabs which he had observed in the West Indies’. In fact, Thompson used his observations of land crabs to support his theory, point- ing out that even such land dwellers must lay their eggs in the sea and give their young a perilous life there, so that they may undergo metamorphosis before going ashore in their adult form. He points out, too, that fifty years earlier, in a work published at Haarlem, Slabber (1778) claimed that he had observed metamorphosis in Zoea. Thompson swiftly disposes of that claim: “The metamorphosis, however, which this observer thought he witnessed is of so different a description that we must either suppose him to have fallen into some error or else there may be Crustacea which pass through some other forms’ He concludes, ‘from much experience’, that Slabber ‘lost his Zoeas in changing the sea water, and that the new form came from the added portion’. Nevertheless, in his own first memoir, Thompson, too, had ‘failed to prove the complete metamorphic cycle because his zoea died in the process of change; it was only by comparing them [sic] with ova from a berried female crab that he was able to deduce the relationship (Wheeler, 1970). What is more, his announcement was accompanied by a second memoir ‘On the genus Mysis (Thompson, 1828), in which he showed that the mysidacean crustaceans hatch in a form very similar to that of the adult and undergo only slight metamorphosis, thus tending to contradict his assertions about metamor- phosis as a characteristic of most crustaceans. And both memoirs appeared only months before the German scientist Rathke, unaware of Thompson’s announcements, pub- lished his own observations on the development of the crayfish (Potamobius, formerly Astacus), now known to be one of the few exceptions to the general rule propounded by Thompson. Rathke’s work demonstrated that the young crayfish hatch at a late stage of development and do not undergo metamorphosis. PROC. LINN. SOC. N.S.W., 111 (2), 1989 52 JOHN VAUGHAN THOMPSON, F.L:S. One might ask why Thompson was in such a hurry to publish when his conclusion was not entirely proved and was supported by only a single observation. The answer lies in the comment made by Stebbing in relation to another of Thompson’s discoveries, the Polyzoa, where an observation made in 1823 was not published for seven years, and so was not credited to him. ‘As we all know, recognition of our discoveries has to date, not from the time when they were made, but from the time when they were published’ (Stebbing, 1910). Thompson knew that others were studying marine invertebrates, and did not want to risk being beaten. As it happened, he was alone in detecting metamor- phosis, and could have afforded to wait. But given Thompson’s reliance on deduction in his first memoir, it is not surprising that many established zoologists treated his claim of metamorphosis in the Crustacea with ‘distinct and often derisory doubt. They resented the overturning of the taxonomy in use at the time, as expounded by Leach and Cuvier. Nevertheless, Thompson con- tinued to announce fresh examples to prove his hypothesis. By the end of the decade, Rathke and others had published admissions that their scepticism had been misplaced and that Thompson was correct (Wheeler, 1975). In the middle of 1830, Thompson’s military career blossomed briefly. On 22 July he was promoted to the rank of Deputy Inspector General of Hospitals (the next step on the promotion ladder for surgeons) as a reward for his long and zealous service. The pro- motion promised him an income sufficient both to support his family in the style he thought fitting and to continue publishing the results of his research. Then, as suddenly as his fortunes rose, they fell (Johnston, 1917). On 9th December 1830 he was retired to half pay, along with 13 other officers of his rank who experienced the same meteoric rise and fall. At age 51, with a wife and six children (under fifteen?), he saw his income drop from 30 shillings a day to 17. Not only did the reduction threaten the prospects of his family, but it also made it impossible for him to go on publishing his Zoological Researches without help. Some might think he should have been able to bring up a family on £300 a year. Thompson did not. Consequently, the proceedings of a meeting of the Zoological Society of London a month later, on 11 January 1831, (Zool. Soc., 1831), show that it received ‘an Address by Mr J. V. Thompson, Esq., “Io the members of the Zoological Society, and the Zoologists of the United Kingdom in general”, soliciting such support, by subscription, as may enable him to continue, without further loss, his Zoological Researches and Illustrations. This address is printed, together with a list of the subjects of some of the preceding memotrs, on the cover of the fourth number of the Researches, which was at the same time laid on the table’. Stebbing, writing eighty years later, referred to ‘a pathetic appeal to the scien- tific world to furnish him with 150 subscribers, as his private income would no longer bear the sacrifice till then entailed by the publication of his researches’ (Stebbing, 1910). Thompson’s financial setback did not prevent him from announcing further evidence of metamorphosis in the Crustacea, in response to a complaint from Vigors (1830) in the Zoological Journal that his conclusions had been too sweeping. His letter to the Secretary of the Zoological Society dated ‘Cork, Dec. 16, 1830 is reported thus (Zool. Soc., 1830: internal quotations are from Thompson): ‘In it, Mr Thompson urges, in support of the universality of a metamorphosis among the C7ustacea, that he has ascer- tained the newly hatched animal to be a Zoea in eight genera of the Brachyura, viz. Cancer, Carcinus, Portunus, Eriphia, Gecarcinua, Thelphusa?, Pinnotheres, and Inachus; and in seven Macrourous genera, viz. Pagurus, . . . and Astacus. “These embrace all our most familiar native genera of the Decapoda”. The Lobster, or Astacus Marinus, Mr Thompson states, “does actually undergo a metamorphosis, but less in degree than in any of the other enumerated genera, in its first stage being what I should call a modified Zoea . . .; in short, such an animal as would never be considered what it really is, was it not obtained PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 35) by hatching the spawn of the Lobster”. In the other indigenous species of Astacus, the River Crawfish, it would appear from the excellent treatise of M. Rathke on the de- velopement [sic] of its eggs, that the young are hatched in a form according with that of the fully grown animal. Mr Thompson, however, suspects that some source of error may exist in these observations. “If it should be found otherwise, it can only be regarded as one solitary exception to the generality of metamorphoses, and will render it necessary to consider these two animals for the future as the types of two distinct genera”. . .’ This letter is an interesting demonstration of Thompson’s confidence in the accuracy of his observations: he did not hesitate to question the opinions and obser- vations of established and respected naturalists when they conflicted with his own. The Cirripedes Thompson’s third important achievement in marine biology was his discovery that cirripeds are Crustacea: in the system proposed by Cuvier, they had been designated as a class of the Mollusca. He published his discovery in the third number of Zoological Researches, Memoir iv: On the Cirripedes or Barnacles; demonstrating their deceptive character, the extraordinary Metamorphosis they undergo, and the Class of Animals to which they indisputably belong (Thompson, 1830a). Wheeler (1975: internal quotation is from Thompson and Darwin) explains the significance of the discovery thus: ‘Again this presented a revolutionary concept as far as the higher classification of barnacles was concerned. Among the barnacles two major groups are well known, the goose or ship’s barnacles, which look somewhat like small, light coloured mussels on a stalk, and the acorn barnacles, which are so abundant on rock and timber on shore. Earlier naturalists had been much exercised about the true relationships of these animals. “Thompson stumbled on the correct solution to their relationship by, as he put it, “the result of chance rather than of design and industry”. In 1823, crossing on the ferry at Passage Cove, he trailed his small muslin net behind the boat to catch a quantity of marine zooplankton. Among them were translucent elliptical animals a tenth of an inch in length, evidently crustaceans but of a form undescribed by earlier naturalists. On 1 May 1826 he collected further specimens, and in the light of his knowledge that larval decapod crustaceans were entirely dissimilar to the adults kept them in order to observe any metamorphosis which might take place. He had the satisfaction on 8 May of seeing some of these elliptical animals metamorphose, and found them firmly attached to the bottom of the glass container, perfectly recognizable as young acorn barnacles. On 10 May another of these larvae was seen to cast its outer skin and settle to the bottom as a barnacle. In 1835 Thompson described the larvae of the goose barnacle, which he found on a ship’s bottom in 1830. ‘Again, though Thompson had firmly placed the barnacles within the Crustacea and showed that they too enjoyed a planktonic larval stage before settling, established zoologists were slow to accept his findings. However, Darwin (1851) in his monograph on the group recognized Thompson’s contribution as a “capital discovery” .’ Of course, Thompson would not have ‘stumbled on the correct solution had he not been looking with the eyes of one who knew that metamorphoses occurred in Crustacea. The Royal Society’s Catalogue of Scientific Papers 1800-1863 (Royal Society, 1871) records some of the reprints of Thompson’s papers which had appeared by 1831 in English and French journals. An illustration of the way Thompson’s name cropped up incidentally in scientific discussions in the early 1830s comes from the proceedings of the Zoological Society on 11 January 1831 (Zool. Soc., 1831). A question was raised at this meeting about the cause of phosphorescence in sea water. ‘It was remarked that Commerson and others have PROG. LINN. SOG. N.S.W., 111 (2), 1989 54 JOHN VAUGHAN THOMPSON, F.L:S. attributed the phenomenon described to putrefaction of animal matters, . . . Sir Joseph Banks, Dr Macartney, and others, on the contrary, have referred it to the presence of marine animals, principally Crustacea; and the existence of such, as the cause of this appearance, has been recently insisted on by Mr J. V. Thompson,’ Polyzoa The last of Thompson’s four important discoveries was that of Polyzoa, a new Animal discovered as an Inhabitant of some zoophytes (Thompson, 1830b). He showed that the term zoophytes had been used to cover a mixture of animals superficially alike but essentially different in structure. Relying on Wheeler again: ‘[ Polyzoa| had been formerly included as part of a heterogeneous collection of enigmatic invertebrates, the so-called zoophytes; but he showed that they were distinct from the colonial hydroids and the ascidians, with which they had been sometimes confused. The term Polyzoa received considerable usage, especially in Great Britain, but it was eventually dropped in favour of Bryozoa, which had been proposed almost contemporaneously’ (Wheeler, 1970; supported by Stebbing, 1910). Because of Thompson’s preoccupation with natural history, it is not surprising to find that he left only one paper on medical science. Although he was a prolific writer and despite more than forty years in the Army Medical Department, his only recorded pub- lication in the field is an 1832 pamphlet entitled The Pestelential Cholera Ummasked . . .” Wheeler (1970) describes it as ‘a work devoted to diagnosis and treatment of cholera, but exhibiting little understanding of the causative factors involved in the disease’. Perhaps the critic expected too much. At the time, no one else understood such matters, either. In fact, the pamphlet conveyed the prevailing wisdom about the disease and it was topical: cholera had been spreading westward from Asia during the 1820s, and the first major outbreak in the British Isles occurred in 1832. Wheeler (1975) reports in passing that in 1833 Thompson's collection of in- vertebrates was purchased by the Royal College of Surgeons, Dublin. Unfortunately, the collection has not survived. In the light of his financial setback at the end of 1830, one wonders what domestic crisis forced Thompson to sell it. SCIENTIFIC DISPUTES AND FINANCIAL DIFFICULTIES Meanwhile, Thompson’s conflict with the zoological establishment in London became more and more acrimonious as his new concepts threatened to overturn fundamental systematics. As Wheeler (1970) says, ‘His correspondence and published writings of the time suggest a man impatient with the conservatism of his opponents, and eventually embittered by their opposition . . .’ Elsewhere, Wheeler (1975) says: ‘His tendency to write in a forthright style cannot have endeared him to his opponents’. Meanwhile, too, Thompson’s money problems became more and more embarrass- ing. On 8 May 1835 he informed the Treasurer of the Linnean Society that he was ‘unable to send the large amount due’ and ‘craved the indulgence . . . in the hands of the Committee of remitting the payments in special cases of disability. He pointed out that he had not been attending meetings or receiving copies of the Society’s ‘valuable Trans- actions’ while his fees (at the time £3 per annum) were in arrears. He explained that he was claiming the indulgence ‘. . . on account of the pecuniary loss sustained by the pub- lication of my Zoological Researches and the great expense incurred by the prosecution of those discoveries by being obliged to keep up an establishment on the sea-side during a great part of the period, & the disbursements consequent on boat hire travelling back and forward, &c (Linn. Soc., 1835). According to its minutes for 22 May 1835, Council ‘resolved that the request be PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 55 acceded to on account of his high merit as a naturalist and the papers he had supplied for the Transactions of the Society. In the middle of dealing with both his financial difficulties and a bitter scientific controversy, Thompson was removed from the scene. Colonial authorities in London had for some time been anxious to reduce the cost of the separated convict medical ser- vices and military medical services in New South Wales and Van Diemens Land. Early in 1835, the Government decided to combine both services in each colony under the supervision of senior Army surgeons (Colonial Office, 1834). Accordingly, in June 1835, John Vaughan Thompson was offered an appointment as Deputy Inspector-General of Hospitals in New South Wales (Colonial Office, 1835a). Distressed as he was, he could not refuse the offer, since it meant he would be restored to full pay from 7 August 1835 (Johnston, 1917). Only money could have taken him so far abroad; and it could not have done so at a worse time, because scepticism about his Zoological Researches was at its height. In the very week that Thompson was offered his new appointment, his most implacable opponent, J. O. Westwood, F.L.S., secretary of the Entomological Society and through that office committed to retaining the old taxonomy, attacked Thompson's work by presenting a long paper to the Royal Society On the supposed existence of Metamorphoses in the Crustacea. Westwood (1835) used special cases and the opinions of a number of respected authori- ties to deny outright the evidence that Thompson presented. He was quite wrong, but the error did not make his criticism any less credible to many of his listeners and readers, who shared his misconceptions. Patronizingly, he pointed out that ‘the accuracy of [Thompson's] beautiful figures deserves the highest praise’ and summed up by saying, ‘Although disagreeing with Mr Thompson in respect to his theory, I have already stated that his figures are very faithful delineations of nature’. In all else, he set out to discredit Thompson. Perhaps that paper spurred others, some of them also severe critics, to take Thompson seriously and make their own observations. Nevertheless, it took some time to verify Thompson's work, and it was not until the end of the decade that impartial zool- ogists repeated his observations and admitted that he was right (Wheeler, 1975). Not that Thompson himself had slackened in his efforts to substantiate his dis- coveries. He had been continuing his research and his writing, and a number of his papers appeared in various journals in 1835 and 1836 (Royal Society, 1871). Some of them appeared even in the heart of the enemy’s territory, the Entomological Magazine (Royal Society, 1871). DEPUTY INSPECTOR-GENERAL OF HOSPITALS, NEW SOUTH WALES, 1836-1844 Surgeon-Superintendent of the ‘Boadicea’ In June 1835, however, Thompson had the more pressing problem of arranging passage to New South Wales for himself, his wife and four children. In his fifty-sixth year he was about to begin a new career in a new land, and turn his back on the researches which had occupied him for the past 20 years. By the beginning of August 1835 he had settled most of his affairs in Ireland and returned to London. He left unpaid a seemingly trivial account for goods to the value of £6.10.7% with Mrs Dwyer of 89 Grand Parade, Cork, which was to haunt him for years (Colonial Office, 1841a). On 7 August 1835, the day he returned to full pay, he wrote a memorial to the Secretary of State for the Colonies, seeking a grant of land in New South Wales as a recompense for losses he and his father had suffered in government PROC. LINN. SOC. N.S.W., 111 (2), 1989 56 JOHN VAUGHAN THOMPSON, E.L:S. service (Thompson, 1835)*. The Secretary of State replied that the regulations did not allow him to make the grant (Colonial Office, 1835b). To minimize his expenses, Thompson secured the post of Surgeon-Superintendent on the emigrant ship Boadicea taking 200 unmarried females and about 60 people in family groups to Van Diemens Land. For this service he received £50 and his own free passage, but he had to pay what seems an exorbitant price of £159 for cabin passages for his wife and children. On 8 August 1835 he signed his contract with the Secretary of the Emigration Committee, undertaking to be available from 26 September ‘to inspect the emigrants and witness their promissory notes’ (Colonial Office, 1835c). The Boadicea sailed on 1 October 1835, and arrived in Hobart Town on 4 February 1836. It is an extraordinary coincidence that Charles Darwin, who would later praise Thompson’s work on barnacles, sailed into the Derwent River next day on the Beagle. There is no evidence that Thompson and Darwin met, although they were in Hobart Town at the same time for more than a fortnight. There were no deaths on the 128-day voyage. The Surgeon-Superintendent reported that: ‘I am quite confident that it was to [the] regulation of their diet, the with- drawal of their wine while in the Torrid Zone and the promptitude with which every case of sickness was met that under Divine Providence I am indebted for the preservation of very many lives — not that we escaped our share of sickness nor the occurrence of several very hopeless cases & almost miraculous escapes} (Colonial Office, 1836b). After almost a month waiting in Hobart Town, Thompson and his family secured a passage to Sydney on the North Briton, arriving on 15 March 1836 (Herald, 1836). He took up his new duties in the Office of Deputy Inspector General of the Convict [Civil] and Military Hospitals in the Colony of New South Wales on 1 April. Sir James McGrigor, head of the Army Medical Department since 1815, was well aware of the qualities of his senior officers, and proved a good friend to Thompson in all that was to follow. He had sent Thompson his instructions on 20 August 1835 (Colonial Office, 1835d). The Deputy Inspector General’s mission was to manage Convict and Military Hospitals as a single service, as far as possible under the Hospital Regulations of the Army. He was to give particular attention to keeping down costs, to controlling the requisition and issuing of medicines and making regular returns about their use, and to regulating the access by civil servants and other free residents to hospital services. With uncanny prescience, McGrigor (Colonial Office, 1835d) advised Thompson against possible pitfalls: ‘In the allotment of the duties civil and military you will take care as much as possible to preserve the harmony necessary to good order; and in assimilating the duties you will not unnecessarily disturb the Colonial Surgeons’ appointments, and on all occasions obtain the approbation of the Governor and proper authorities in recommending any change of duty among inferior officers and servants’. Early Administrative Problems In ‘preserving harmony’ and ‘not disturbing the Colonial Surgeons, Thompson failed utterly. Accordingly, his previous biographers have either discreetly ignored the New South Wales part of his career or written about that to the exclusion of all else. Part of the reason for his failure was undoubtedly the forthright and abrasive per- sonality that he had already revealed in his scientific disputes and in his rejection of an offensive complaint about rations from the sea-lawyers on the Boadicea (Colonial Office, * This memorial has been mentioned already as the source of hitherto unknown information about Thompson’s family and his service in Mauritius. { Im fact, it is the frankness and directness of Thompson’s reports on the emigrants that first attracted the writer’s attention to him. He seemed so determined to face opposition head on, and to do it in writing, (Campbell, 1988) that I was compelled to find out what happened to him afterwards — and before. PROG. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL D// 1836d). In 1838, Governor Gipps said of him: ‘Dr Thompson, whatever may be his other merits, wag not the man calculated to carry successfully into effect a measure In itself un- palatable to his subordinates, being on the one hand wanting in blandness of manner or conciliatory address, before which opposition might have gradually given way, and on the other in that firmness and decision of purpose which would have overruled it’ (Colonial Office, 1838a). But as MeGrigor wrote: “His situation was no easy one for any man of common patience and temper. It was perhaps to be expected that an officer experienced in the duties, sent out to control expenditure, diminish emoluments and supervise the manner in which the several officers discharged their duties would be unacceptable, let his address and manner be what they would’ (Colonial Office, 1839). First, he had to serve three masters: the Governor and Secretary of State for the Colonies in matters relating to the Convict [Civil] establishment, the Secretary at War in matters relating to the Army, and the Director General of the Army Medical Service in regard to professional reports and returns. What is more, running through his deal- ings with all three was an undercurrent of resentment from officials in the Colonial Office about the intrusion of the Army Medical Service and the War Office into their domain. Hence, the Governor and his London superiors seemed to take some pleasure in pointing out Thompson's failings to McGrigor, who was responsible for appointing him and for disciplining him thereafter. Second, through an incredible bureaucratic bungle, he arrived in Sydney to find that Dr James Bowman, who had joined the colonial medical service in 1817 and had been its head since 1819, was still in that position, and no arrangements had been made to remove him from it. Consequently, one of the first recommendations Thompson had to make was that his predecessor should be stood down. In the event, it took two years for the despatches necessary to terminate Bowman’s appointment to pass back and forth between Sydney and London. Meanwhile, Bowman remained on strength without duties, devoting himself to his extensive private practice and the pastoral schemes of his father-in-law, John Macarthur. Even without those supplements, his unearned salary was some £200 greater than the pay for Thompson’s active Army rank. Third, the other colonial surgeons, especially James Mitchell, who had been on good terms with Bowman, also resented ‘Thompson’s arrival. His lack of a medical degree did not go unnoticed, and the colonial surgeons were aware of his proposals to replace them by army surgeons at lower rates of pay when their posts became vacant. The security of their lucrative private practices was also threatened by their new lability to transfer. And Thompson was less than tactful towards them. He was critical, in reports to Sir James McGrigor, of their private practices, their drinking, and their failure to cooperate with him. Fourth, to cap Thompson's difficulties, the Governor reported at the end of 1836 that ‘Mr Thompson has not as yet presented any distinct and comprehensive arrange- ment for. . . hospitals in this Colony, chiefly | apprehend from the infirm state of health under which he has for some time been suffering (Colonial Office, 1836c). In fact, Thompson had devoted a good deal of time to pointing out that his travel- ling allowance did not cover the cost of the journeys he was required to make; comparing his own pay and allowances with those of other senior officials; confronting the colonial surgeons; and complaining to London about the obstacles the latter were putting in his path. ‘To put the matter bluntly, the duties Thompson was expected to carry out, and the circumstances in which he found himself, were beyond the powers of an elderly, ailing, quarrelsome and forthright army surgeon who had been five years retired, and who in PROG. LINN. SOG. N.S.W., 111 (2), 1989 58 JOHN VAUGHAN THOMPSON, F.L.S. any case would rather have been working at his microscope. The consequences were inevitable. Nevertheless, Sir James McGrigor in London argued in his support: ‘Mr Thompson complains that he has met with vexatious obstruction from some of the Colonial Medical Officers at the outset; perhaps it 1s but natural that some of these gentlemen should, at first, feel a new control irksome, and complain of more duty being imposed on them than under the old system so as to interfere with lucrative private practice or their avocations as agriculturalists . . . ‘I am not prepared in every instance to support the tone of Mr Thompson's querulous correspondence, and I have so expressed my disapprobation of it that I trust this zealous and experienced officer . . . will henceforth be most respectful in all his cor- respondence . . . and I am of the opinion that henceforth no infirmity of temper will be betrayed by the Deputy Inspector General of Hospitals. ‘In justice, however, to Mr Thompson I ought to say that during a long period of service I have ever found him an able and honourable officer, one who has constantly evinced the most indefatigable zeal on every service on which he has been employed, acting with discretion, prudence and forbearance’ (Colonial Office, 1837a). Despite this testimonial and promise for the future, in September 1837 a dispute known as ‘the Mitchell case’ made public a conflict which had been festering since Thompson’s arrival and which dragged on for almost four years. Briefly, Mitchell, who had been in rancorous disagreement with Thompson from the outset, claimed to have misunderstood an instruction, and so failed to attend as medical officer at a flogging. Consequently the Governor dismissed him for disobeying an order. In response to pub- lic support for Mitchell, Thompson inflamed the debate by writing a letter to a newspaper; in it he referred selectively to some comments made about Mitchell by the Governor. It is true that the Governor had more than once criticized the way the colonial surgeons, including Mitchell, responded to Thompson’s instructions; but he had criti- cized the tone and nature of the instructions themselves almost as often. Mitchell sued Thompson for libel, and won. In the meantime, he had appealed against his dismissal. The appeal led eventually to an inquiry by Governor Gipps: as a result, Mitchell was reinstated in 1841 for one day, for the sole purpose of allowing him to retire without a stain on his name. Although the finding supported Thompson’s rank and office, he suffered more than Mitchell, who in fact had little to lose. But Thompson was left a broken man, after yet another acrimonious conflict which had simply deepened his bitterness and disillusionment. An account of the quarrel between Thompson and Mitchell, distinctly unsym- pathetic to Thompson, is provided by McIntosh (1956). It contains a good deal of accu- rate detail but does not, in the writer’s opinion, sufficiently recognize the problems Thompson faced in dealing with an entrenched Sydney establishment. A good summary of the evidence is given in Gipps’ Report on his Review of the case (Colonial Office, 1841b). Thompson made other administrative mistakes, too. Soon after his arrival, he was taken to task for writing personal letters to the Secretary of State for the Colonies about mismanagement by certain officers of the colony who were outside his jurisdiction. He was told to stop his ‘political investigations and writings’ (Colonial Office, 1837b), and concentrate on his public duties. And as time went on, he displayed an increasing reluc- tance to provide the reports and accountings required of him, leading to comments about his ‘inflexible taciturnity’ and ‘habitual delay’ (Colonial Office, 1842a). What is more, he did not bring about the hoped-for reduction in costs. PROG. LINN. SOG. N.S.W., 111 (2), 1989 HUGH CAMPBELL 59 Sustained Interest in Natural History Despite Thompson’s difficulties with his official duties in New South Wales, he continued to show his interest in natural history. Except for the dedication of his Zoologi- cal Researches to Sir James McGrigor, he had scrupulously kept this interest separate from his life as an army surgeon. He began optimistically. On 20 April 1836, he wrote to his old acquaintance, Alexander Macleay, the Colonial Secretary, expressing his disappointment at not find- ing ‘the Society”™* flourishing, and his hope that Macleay would re-establish it and name some original members. He also made a number of suggestions about the scope and purpose of a natural history society, recommending a wide membership (Thompson, 1836). It appears that this letter was Thompson’s acceptance of an invitation to join ‘A Committee of Superintendence of the Australian Museum and Botanical Garden’ (Gazette, 1836). Macleay became chairman, and Thompson was a member. The minute books of the Committee show that he attended meetings regularly from 1836 to 1843 and was a member of the Sub-Committees for both the Botanical Gardens and the Australian Museum (Australian Museum, 1836.) Meanwhile, he still found time to look to his scientific reputation and contacts. His letter to Julien Desjardins, President of the Natural History Society, Mauritius, claim- ing authorship of the Catalogue of Exotic Plants in Mauritius has been mentioned already. A letter from the Curator of the Australian Museum records that ten specimens of native birds were given to J. V. Thompson ‘for the Mauritius’ in 1837 (Australian Museum, 1837). He is also on record in 1839 as having sent cotton seeds to Mauritius in 1838; and the 1840 Annual Report of the Mauritius Natural History Society records that “Thomp- son is the only one of our New South Wales correspondents who has not given us up’ (Vaughan, 1953). Thompson would have been heartened, too, to find that in 1839 Du Cane, a friend of Macleay, was one of those who had confirmed his observations on the metamorphosis of the Crustacea (Fletcher, 1920). Removal from Office Ever since Thompson had left Cork in 1835 without settling his account with Mrs Dwyer, she had been trying to get her money. Failing in appeals directly to Thompson, she had then unsuccessfully sought to have Sir James McGrigor order Thompson to pay. Getting no satisfaction in that quarter either, on 28 October 1841 she complained to the Secretary of State for the Colonies. It was Thompson’s bad luck that her letter arrived only a fortnight after the Secretary of State had been considering Governor Gipps’s report on the Mitchell case, and so Thompson's name was fresh in his mind. There is no record to show that Thompson paid the bill, in spite of a fresh instruction to do so. During 1842, Thompson’s neglect in submitting reports and returns became intolerable to his superiors (Colonial Office, 1842b). Finally, in November, he was given six weeks to submit a particular return. When he did not meet the deadline, preliminary steps were taken to replace him (Colonial Office, 1843a). The Governor reported to the Secretary of State, who in his turn asked McGrigor to remove Thompson. By this time it was July 1843. In the letter to McGrigor, the Colonial Office reminded him of Mrs Dwyer’s case, saying that ‘this renewed proof of that officer’s negligence’ seemed to call for his suspension from duty (Colonial Office, 1843b). McGrigor replied that he was already aware of the situation, and recognized the need to act: ‘.. . for some time back I have had occasion to notice to Mr Thompson many instances of neglect in conducting the public duties and of his rarely performing the duty of inspecting the hospital, but finding this proceeded from greatly impaired * Presumably he was referring to the defunct Philosophical Society of Australasia. PROC. LINN. SOC. N.S.W., 111 (2), 1989 60 JOHN VAUGHAN THOMPSON, FL.S. health, with mental as well as bodily infirmity, I informed him of my intention. . . of his being placed on the half pay of his rank in the army’ (Colonial Office, 1843c). McGrigor followed up his official letter with a private one to Stephen, his counter- part in the Colonial Office, presumably to make sure that no one proceeded with the idea of suspending Thompson or dismissing him outright, and so cutting off his income: ‘The neglect of his duties by Mr Thompson .. . is not unknown to me. I regret to say that for a considerable time back I have had cause to complain of the imperfect manner in which he has performed his duties. I have reason to believe that this has proceeded from the broken down health of Mr Thompson who is no longer equal to the multi- farious duties of the responsible situation in which he is placed’ (Colonial Office, 1843d). It is curious that Governor Gipps made no mention of Thompson's deteriorating health, which had been mentioned by Bourke at the end of 1836 but ignored thereafter in despatches from New South Wales. Hence one wonders how McGrigor came to his conclusion that ill health was an important reason for Thompson’s poor performance. But the question is academic: Gipps had secured Thompson’s removal. Thompson was allowed to remain in office until his successor arrived in New South Wales (Colonial Office, 1843e). He relinquished his position on 1 February 1844, and his retirement to half pay was gazetted on 11 February (Johnston, 1917). It should be said in his defence that, despite the problems described above, he sur- vived for almost eight years in a position which had been almost untenable from the out- set — more than twice as long as his counterpart in Van Diemens Land, who had taken up duty at the same time and was superseded in 1839. And no one seems to have com- plained about the quality of the medical service under his regime. In fact, at the end of 1839 Gipps had made a special point of remarking that he had no complaint about the way the ‘strictly professional duties of the Medical Department in this Colony are performed’ (Colonial Office, 1838b). Thompson's ‘taciturnity’ and failing health in the 1840s did not prevent him from returning to his old interest in botany, and four more papers from him on the cultivation of cotton and sugar cane were published in the /ndian Journal of Agriculture between 1842 and 1845 (Royal Society, 1871). When Thompson retired, one piece of business with the Colonial Office was still unfinished. On 6 April 1843, having heard of a plan for sending young prisoners from Parkhurst to the colonies for rehabilitation, Thompson wrote another characteristically naive proposal to the Secretary of State for the Colonies. He announced that he had ‘just purchased a farm (part of Eagle Farm) about seven miles from Brisbane ‘Town, and [had] at present about 200 acres of clear tillage land’. His scheme was hastily conceived and impracticable. He said he would be ‘most happy to try the experiment with from 10 to 20 of the Parkhurst boys . . . [T]hey will be removed from all temptation to steal or pick pockets and will be employed in gardening, assisting the agricultural labourers and vine dressers and in looking after the farm work. My object is to give a fair trial to tropi- cal agriculture and in particular to cotton, sugar cane and coffee in addition to the vine and other colonial productions’ (Thompson, 1843). Perhaps Thompson wanted only to tell someone that he had just bought a property to replace the land his father had lost sixty-five years earlier. Whatever he intended, his plan misfired. It was typical of his luck that his application for Parkhurst boys was con- sidered and rejected just the day before the Secretary of State approved his transfer to half pay. His Lordship remembered the letter, and saw it as just one more proof of Thompson’s inattention to his real duties (Colonial Office, 1843f). Thompson’s last official letter is a sad reminder that mail took several months to pass between London and Sydney. A week after Thompson had been retired to half pay, the Governor called for a copy of his letter about the Parkhurst boys. Jno V. Thompson, PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 61 Deputy Inspector General, replied on 21 February 1844: ‘I have the honor to acknowl- edge receipt of your communication of the 17th instant (44/14), and regret that I am incapable of supplying the copy of the letter addressed to the Secretary of State of 6th April last, considering it to be a private affair and only a request, which, having been deprived of every means of accepting now by the loss of my appointment, I must of necessity relinquish (Thompson, 1844). As far as is known, Thompson remained in Sydney for the next three years, and so did not have a chance to occupy his land at Moreton Bay. He died at his residence in Liverpool Street on 21 January 1847, and was buried in the Parish of St Lawrence, County of Cumberland (Registrar General, 1847). Presumably his grave was in the old Devonshire Street cemetery, now the site of Central Railway Station. He directed that the monies from his estate should be used in the first instance to pay his debts, and that what remained should be paid to his ‘dear wife for her use entirely (Supreme Court, 1847). CONCLUSION Since his death, John Vaughan Thompson’s contribution to natural science has been recognized spasmodically. A crustacean, Vaunthompsonia, was named after him, as was a plant from Madagascar, Dezdamia thompsoniana, though the latter was subsequently given another name. He merits a line in some histories of zoology (Singer, 1959); Lankester (1890) gives him a full paragraph in the Zoology article in The Advancement of Science, remarking that “Thompson made ... great discoveries, which seem to have fallen in his way in the most simple and natural manner, but must be regarded really as the outcome of extraordinary genius’; and he 1s the subject of entries in the Dictionary of National Biography and the Dictionary of Scientific Biography under his own name. His Zoo- logical Researches were reissued in a facsimile edition in 1968. It is interesting that the burning issues of his day were not seen by later biographers to be controversial. For instance, metamorphosis in Crustacea, the subject of acrimonious debate in the 1830s, was overlooked by Lankester, and hardly mentioned by Stebbing (1910) in the paper he wrote to mark the centenary of Thompson’s election to a Fellowship in the Linnean Society. Today, however, 140 years after his death, the way he approached natural history is more important than the truths he revealed. He was one of the genuine discoverers in the age when gentleman naturalists played a significant part in scientific inquiry, and even in that he illustrates a point: he shows how hard it was for a man short of money, and working alone, to have his work noticed. Previous biographers have praised his per- ceptiveness, and shown how his discoveries resulted from acute practical observation. But they have not made enough of his persistence, his solitariness and the disappoint- ments he suffered. He was a ‘difficult’? man and yet he was able to make a notable contribution to both botany and zoology. And that is only one side of his life. It is too easy to forget that his eight-year stint in New South Wales, where the odds were against him from the first, was the only blemish on a career in the Army Medical Service spanning forty-five years. On balance, his unhappy inability to get on with people neither cancels out his work as a surgeon nor dulls the brilliance of his lonely scientific achievements. It is a pity that he was not in England when his discoveries were finally recognized. There he might have been honoured instead of being ignored. The careful obituary in the Sydney Herald of 26 January 1847, with no mention of friends and colleagues who appreciated him, emphasizes his loneliness. And it was John Vaughan Thompson’s usual luck that the age was wrong. He was in his 68th year, not his 63rd:‘. . . in his 63rd PROG. LINN. SOG. N.S.W., 111 (2), 1989 62 JOHN VAUGHAN THOMPSON, EL.S. year, after long-continued illness — distinguished for his achievements in zoology and botany — possessing talents of no common order — and estimable in every relative duty of life — he is deeply lamented by his afflicted family, to whom his loss is irreparable’ (Herald, 1847). ACKNOWLEDGEMENTS The Public Records Office, London, has granted permission for reproduction of the Crown Copyright Colonial Office documents quoted in this paper. Acknowledge- ment is made to the Australian Joint Copying Project of the National Library of Aus- tralia and the Library of New South Wales for the microfilm of some of those documents, and to the Archives Office of Tasmania and the State Library of Tasmania for the remainder of the documents and for the facilities for reading all of them; to the Archives Office of New South Wales, the Library of the Australian Museum, the Library of the University of Edinburgh and the Library of the Linnean Society of London for the references ascribed to them below; and to the Archives Office of the Government of Mauritius. Thanks are due to Guy Rouillard, President of the Historical Society of Mauritius, Gina Douglas, Librarian and Archivist of the Linnean Society of London, P. H. Sooprayen, Chief Archivist, Government of Mauritius, Jo Currie, Library Assistant, Special Collections, University of Edinburgh Library, and Mary Kumvaj, Deputy Librarian, Australian Museum. Each took the trouble to send me more information than I asked for. References Key to abbreviations used in the text: ADB = Australian Dictionary of Biography, 1967; DNB = Dictionary of National Biography, 1900; EB = Encyclopedia Britannica, 1968; Linn. Soc. = Linnean Society of London; Zool. Soc. = Zoological Society of London. Key to abbreviations used in the references: CO xx/xx = Colonial Office Item Numbers; AJCP xx = Numbers of Australian Joint Copying Project Microfilm Reels; HRA = Historical Records of Aus- tralia volumes. Otherwise standard abbreviations are used for societies and journals. Since many of the references are to manuscript letters and other records, the references are presented in two parts: Part 1: Published Works and Part 2: Unpublished letters, etc. Part 1: References to Published Works ADB, 1967. — Australian Dictionary of Biography. D. PIKE (ed.). Melbourne: Melbourne University Press. BARNWELL, P. J., 1941. — Thompson, John Vaughan: Dictionary of Mauritian Biography, No. 3: 94-95. BRITTEN, J., 1912. — John Vaughan Thompson. /. Bot. Lond. 50: 169-171. CAMPBELL, H., 1988. — A proper class of female emigrants. Zasmanian Historical Research Association Papers and Proceedings. Vol. 35, No. 2. A detailed account of the voyage of the Emigrant Ship Boadicea, J. V. Thompson Surgeon-Superintendent. DARWIN, C. R., 1851 — A Monograph on the Sub-class Cirripedia with figures of all the species. Lepadidae: 8. London: Roy Society. DNB, 1900a. — Vaughan, John: Dictzonary of National Biography, 58: 169. London: Oxford University Press. ——., 1900b. — Thompson, John Vaughan: Dictionary of National Biography, 56: 218. London: Oxford Univer- sity Press. EB, 1968. — Monmouth Court House, Battle of: Encyclopedia Britannica. Chicago: Encyclopedia Britannica Inc. ENCYCLOPEDIA AMERICANA, 1983. — Havana: Encyclopedia Americana. Danbury, Conn.: Grolier Inc. FLETCHER, J. J., 1920. — The Society’s heritage from the Macleays. Proc. Linn. Soc. N.S.W. 45: 626-627. GAZETTE DE LISLE Maurice, 1816. — Projected departures from Port Louis. Samedi 8 juin 1816. Mauritius. Tome 2, page 2, cols. 3 & 4. GAZETTE, 1836. — New South Wales Government Gazette. No. 226, 14 June 1836: 451. HERALD, 1836. — Shipping Arrivals. Herald, Monday 21 March 1836, page 2. Sydney. ——,, 1847 — Obituary. Herald, 26 January 1847, page 3. Sydney. PROC. LINN. SOC. N.S.W., 111 (2), 1989 HUGH CAMPBELL 63 INTERNATIONAL GENEALOGICAL INDEX, 1981. — Index for Northumberland, C0664. International Genea- logical Index. Salt Lake City: Church of the Latter Day Saints. JOHNSTON, WILLIAM, 1917. — Roll of Commissioned Officers in the Medical Service of the British Army, 1727-1896, H. A. L. HOWELL, (ed.): 132. Aberdeen: University Press. LANKESTER, E. Ray, 1890. — The history and scope of Zoology. The Advancement of Science, Occasional Essays and Addresses: 335-336. London: Macmillan. MANNICK, A. R., 1979. — Mauritius, The Development of a Plural Society: 39. Nottingham: Spokesman. McIntosh, A. M., 1956. — The case of Dr James Mitchell. Med. J. Aust. (5) August 4, 1956: 161-168. ROYAL SOCIETY OF LONDON, 1871. — Thompson, John Vaughan. Catalogue of Scientific Papers 1860-1863, 5: 958-959. (Metuchen, N.J.: Scarecrow Reprint Corporation, 1968.) SINGER, C., 1959. — A Short History of Scventific Ideas to 1900: 473. Oxford: Oxford University Press. SLABBER, __, 1778. — Natural Amusements and Microscopical Observations. Haarlem. (Unidentified, apart from mention in J. V. Thompson’s Memoir 1, 1828.) STEBBING, T. R. R., 1910. — On John Vaughan Thompson and his Polyzoa.. . Proc. Linn. Soc. Lond., 123rd session, 1910-11: 64-72. THOMPSON, J. V., 1807a. — Catalogue of Plants growing in the vicinity of Berwick-on-Iweed. London. 1807. octavo, 132 pp. —, 1808. — An Account of some new species of Piper [Pepper], with a few cursory observations on the genus. [2 June 1807]. Trans Linn. Soc. Lond. 9: 200-203. —., 1813. — Description of a new species of the genus Mus, belonging to the section of Pouched Rats. [1812.] Trans Linn. Soc. Lond. 11, part 1 (20 April 1813): 161-163. ——, 1816 (claimed). — A Catalogue of Exotic Plants cultivated in the Mauritius. Mauritius: Baron & Souvignee. ——., 1827. — Memoir on Pentacrinus Europaeus, a recent Species discovered in the Cove of Cork. Cork. 4to, 2 plates. ——., 1828-34. — Zoological Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Cork. (A privately printed collection of six memoirs in five numbers issued between 1828 and 1834. Facsimile published London: Society for the Bibliography of Natural History. 1968. Introduc- tion by Alwyne Wheeler.) —_, 1828. — Memoir J: On the Metamorphoses of the Crustacea, and on Zoea, exposing their singular structure and demonstrating that they are not, as has heen supposed, a peculiar Genus, but the Larva of Crustacea!! Memoir 2: On the Genus Mysis, or Opossum Shrimp. Zoological Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 1. Cork. —, 1829. — Contributions towards the natural history of the Dodo (Didus ineptus, Linn.) a bird which appears to have become extinct towards the end of the seventeenth or the beginning of the eighteenth century. Mag. Nat. Hist. 2: 442-448. ——, 1830a. — Memoir 4: On the Cirripedes or Barnacles . . . and the Class of Animals to which they belong. Zoological Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 3. Cork. ——,, 1830b. — Memoir 5: On Polyzoa, a new animal discovered as an inhabitant of some Zoophites . . . Zoo- logical Researches and Illustrations or, Natural History of Nondescript or Imperfectly Known Animals. Vol. 4. Cork. VAUGHAN, R. E., 1953. — A forgotten work by John Vaughan Thompson Proc. Roy. Soc. Arts G Sez. Mauritius 1, part 3: 241-248. vicors, N. A., 1830. — (In a review of Rathke’s just-published work on the development of the Crayfish). Zool. J. 5, part 18 (June 30, 1830): 241-255. WESTWOOD, J. O., 1835. — Of the supposed existence of Metamorphoses in the Crustacea. Phil. Trans Roy. Soc. Lond. 125: 311-326. WHEELER, A., 1970. — Thompson, John Vaughan: Dictronary of Scientific Biography 13. New York: Scribner. ——,, 1975. — Thompson: Marine Biologist. Brit. Med. J. 3: 534. ZOOL. Soc., 1830. — Minutes, 28 December 1830. Proc. Zool. Soc. 1830: 140. ——,, 1831. — Minutes, 11 Jan. 1831 Proc. Zool. Soc. 1831: 144. Part 2: References to Unpublished Letters and Miscellaneous Records AUSTRALIAN MUSEUM, 1836. — Minute Books of ‘A Committee of Superintendence of the Australian Museum and Botanical Garden’, 1836-1843. Australian Museum Archives. Sydney. ——., 1837. — Records of G. Bennett, Curator, 20 February, 1837. Australian Museum Archives. Sydney. COLONIAL OFFICE, 1834. — CO 202/31 (AJCP Reel 222). Hay to Bourke, 18 December 1834. ——,, 1835a. — CO 201/251 (AJCP Reel 200). War Office to Stewart, Treasury, 15 June 1835. ——., 1835b. — CO 202/31 (AJCP Reel 222). Colonial Office to Thompson, 11 September 1835. —, 1835c. — Archives Office of Tasmania, CSO 1/848/17942. Contract between Marshall and Thompson, 8 August 1835. ——,, 1835d. — CO 201/249 (AJCP Reel 200). McGrigor to Thompson, 20 August 1835. ——,, 1836a. — Archives Office of Tasmania, CSO 1/848/17942. Boadicea Passenger lists. PROC. LINN. SOC. N.S.W., 111 (2), 1989 64 JOHN VAUGHAN THOMPSON, F.L:S. ——., 1836b. — Archives Office of Tasmania, CSO 1/848/17942. Thompson’s Report, 4 February 1836. ——, 1836c. — CO 201/255 (AJCP Reel 203). Bourke to Glenelg, 10 November 1836. ——, 1836d. — Archives Office of Tasmania, CSO 1/848/17492. 14 passengers on the Boadicea to J. V. Thompson, 20 November 1835. ——,, 1837a. — CO 201/269 (AJCP Reel 209). McGrigor to Stephen, 27 June 1837. ——., 1837b. — CO 201/268 (AJCP Reel 212). Thompson to Sec. of State, 23 September 1837. ——., 1838a. — HRA I, Vol. 19; 391. Gipps to Glenelg, 20 April 1838. ——., 1838b. — HRAI, Vol. 19: 396. Gipps to Glenelg, 24 April 1838. ——., 1839. — HRA, Vol. 20: 77. McGrigor to Stephen, 5 February 1839. ——, 1841a. — CO 201/313 (AJCP Reel 336). Mrs Dwyer to Sec. of State for Colonies, 28 October 1841. ——.,, 1841b. — CO 201/306 (AJCP Reel 331). Gipps to Sec. of State, 19 January 1841. ——.,, 1842a. — CO 201/321 (AJCP Reel 340). Gipps to Stanley, 20 June 1842, margin note. ——, 1842b. — CO 201/321 (AJCP Reel 340). Gipps to Stanley, 20 June 1842. ——, 1843a. — CO 201/331 (AJCP Reel 346). Colonial Sec. to Thompson, 7 January 1843. ——.,, 1843b. — CO 201/330 (AJCP Reel 346). Colonial Office to McGrigor, 22 July 1843. ——, 1843c. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 26 July 1843. ——, 1843d. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 1 August 1843. ——, 1843e. — CO 201/338 (AJCP Reel 351). McGrigor to Stephen, 1 September 1843. ——,, 1843f. — CO 201/331 (AJCP Reel 346). Col. Office to Gipps, 7 September 1843. EDINB. UNIv., 1797-8. — General and Matriculation Albums, 1797-8 and 1798-9. Special Collections, University of Edinburgh Library. ——, 1816. — Institutions of Medicine Class Lists, 1816- 7. Special Collections, University of Edinburgh Library. LINNEAN SOCIETY of LONDON, 1807. — Thompson to Seaforth, 7 April 1807. The Linnean Society of London Archives. ——.,, 1809. — Certificate of Recommendation for Fellowship of J. V. Thompson, signed 21 November 1809 by James Sowerby; Alex. Macleay, John Macleay and A. B. Lambert. The Linnean Society of London Archives. ——.,, 1835. — Thompson to Linn. Soc. Treasurer, 8 May 1835. The Linnean Society of London Archives. REGISTRAR GENERAL, N.S.W., 1847. — Deaths, 1847, Vol. 32. Registrar General of New South Wales. Sydney. SUPREME Court, 1847. — Probate, 19 February 1847. Supreme Court of New South Wales. Sydney. THOMPSON, J. V., 1807b. — On the genus Kaempferca. Read to Linn. Soc., 7 April 1807. ——., 1835. — CO 201/251 (AJCP Reel 201). Thompson to Glenelg, Secretary of State for the Colonies, 7 August 1835. —, 1836. — Archives Office of New South Wales, 4.2539.3, file 41/10460. Thompson to Macleay, 20 April 1836. ——, 1843. — CO 201/343 (AJCP Reel 354). Thompson to Sec. of State for Cols., 6 April 1843. ——., 1844. — HRAT, Vol. 23: 439, J. V. Thompson to Colonial Secretary Thomson, 21 February 1844. PROC. LINN. SOC. N.S.W., 111 (2), 1989 A List of the Cryptogams and Gymnospermous Plant Specimens in the British Museum (Natural History) gathered by Robert Brown in Australia 1801-5 E. W. GROVES and D. T. MOORE (Communicated by T. G. VALLANCE) Groves, E. W., & Moore, D. T. A list of the cryptogams and gymnospermous plant specimens in the British Museum (Natural History) gathered by Robert Brown in Australia 1801-5. Proc. Linn. Soc. N.S.W. 111 (2), 1989: 65-102. A list is here published for the first time of the cryptogams and gymnosperms gathered by Robert Brown in Australia and now preserved in the herbarium of the British Museum (Natural History), London. Brief remarks on Brown’s Australian visit and collecting during 1801-5 are given. Correlation with the register of the collection prepared by J. J. Bennett is indicated. E. W. Groves, 143 Westlergh Avenue, Coulsdon, Surrey CR3 3AK United Kingdom (formerly Department of Botany, British Museum (Natural History), and D. T: Moore, Department of Mineralogy, British Museum (Natural History), Cromwetl Road, London SW7 SBD, United Kingdom, manuscript received 12 April 1988, accepted for publication 15 February 1989. . That full account of the Botanical discoveries made during Flinders’s expedition, which the pub- lic had a right to expect, has never appeared...’ J. Lindley, 1844 INTRODUCTION The work of Robert Brown during and after Flinders’ expedition to Australia in 1801-5 is recognized as a significant contribution to Australian natural history in general and Australian botany in particular, but scientific records of the expedition are frag- mentary. Flinders’ (1814) account of the voyage of the Jnvestigator concentrates on geo- graphical, navigational and sailing aspects and is useful for the chronology of the expedition up to his departure in 1803 and subsequent imprisonment at Mauritius. But for botanical details we must rely on Brown. His Prodromus of 1810 was intended to be the main account of the botany, but only one volume of the two proposed was produced, and only a few copies were sold. In due course, this important work was withdrawn. Brown also contributed a modest botanical appendix to the Flinders volumes (1814: 2, 533-613). After the death of the expedition’s commander, Matthew Flinders [1774-1814], British interest in the voyage appears to have declined, and it was not until 1904 that Britten published an account of the plants collected by Brown in Madeira on the out- ward voyage. More recently Rourke (1974) has produced an account of Brown’s activi- ties at the Cape of Good Hope, but there appears to be no list of South African specimens collected by Brown in existence. Brown’s Timor plants are also little known, being only briefly mentioned by Forbes (1885). In Australia Brown prepared lists of plants seen or collected, or both, at some of the anchorages. Where these lists are still extant they remain in manuscript, and as Lindley (1844) pointed out, no published list of Brown's Australian plant specimens was available forty years after Brown returned. It PROG. LINN. SOG. N.S.W., 111 (2), 1989 66 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS was not until the 1870s that a manuscript list of Brown’s specimens was prepared. This is the catalogue referred to below as the ‘Bennett Register’. Recent Australian interest in the voyage began with the account by Burbidge (1956) of Brown’s nomenclatural system (based on Flinders’ chart annotations) for the Aus- tralian collecting localities. A landmark in studies of Brown’s Australian botanical work was reached a few years later when a facsimile edition of the Prodromus (Brown, 1810) was published with an introduction by Stearn (1960a). There has been renewed interest in the natural history aspects of the voyage in the United Kingdom recently with the publi- cation of a paper on the expedition by Edwards (1976) and the publication of the diary of Peter Good [d. 1803] also by Edwards (1981). Some of the paintings by the botanical artist Ferdinand Lukas Bauer [1760-1826] are reproduced by Stearn (1960b), Stearn and Blunt (1976) and Norst (in press). Vallance and Moore (1982) have given an account of the geological aspects of the voyage and Mabberley’s (1985) work on Brown has provided a much-needed biography. Clements (1983) has described the orchids collected by Brown which are in the Lindley Herbarium and is to give an account of those in the British Museum (Natural History) Herbarium. However, there are no published lists of all the 3400 plants collected by Brown in Australia. Further work, which will be published in the near future, includes the edited trans- cription of Brown’s Diary of the voyage (Vallance et a/., in preparation). An important outcome of the Diary publication project is the realization that considerably more speci- mens were collected by Brown than are mentioned in either the Prodromus (Brown, 1810) or the Diary accounts. In his Diary Brown uses a variety of plant names. These include names already published by, for example, Carl Linnaeus [1707-1778], William Aiton [1731-1793] and others. But he also used ‘nicknames’ for species he did not immediately recognize. These are sometimes manuscript names attributable to D. C. Solander [1736- 1786]. Their use is hinted at in a letter (British Library Add. MS 32439, ff61-4 and Fiistorical Records of N.S.W. 4: 776-9) written by Brown to Sir Joseph Banks [1743-1820] from Sydney on 30 May 1802 which contains the following passage: ‘. . . The list. . . is partly compos'd of nicknames given at the moment the plants were collected, and not allow’d since. Some of them, I am afraid, may mislead, and the greater part I am aware are rather barbarous, especially where I have terminated specific names in oides, which I have done when uncertain whether the plant was distinct from a species already known,...’ ‘Oides’ names are mentioned in Brown’s Diary text and as a result of work connected with editing the Diary for publication, the surviving plant specimens gathered by Brown have been located in the BM (NH) Herbarium in order to check if these ‘nicknames’ were mentioned on the original labels. Fortunately most of them are. The locating of Brown’s plants led to the preparation of an index of over 3000 cards (Groves and Moore, 1986), and the cryptogams and gymnospermous representatives located in this way form the basis of this account. ROBERT BROWN’S PLANT COLLECTION A letter from Sir Joseph Banks to Sir John Barrow [1764-1848], written on 9th October 1805 (B.L. Add. MS 32439, f185), reveals that twelve boxes of dried plants and eleven boxes of Bauer’s natural history drawings were landed at Liverpool from the Investigator. Vhere were, in all, 38 cases of natural history specimens (cf. Edwards, 1976). Banks indicates in a letter of 3rd January 1807 (B.L. Add. MS 32439, ff237-245) that about 3600 specimens were in the boxes of plants. Edwards (1976), citing Banks, stated that there were about 700 plant specimens from the south coast of Australia, 500 from the east coast, 500 from the north and 700 from Tasmania; there were some 1000 PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 67 specimens from the neighbourhood of Port Jackson, and 200 from the island of Timor (not discussed here). Brown wrote relatively little on the lower plants, but the existence of his paper on mosses (Brown, 1811) indicates he was to some extent interested in Bryophytes. But his greatest contribution to the plants that are our concern in this paper was the realization that the cycads had coniferous similarities (Brown, 1826). Having survived the voyage to England in a leaking ship, impounding at the Custom House of Liverpool, and a journey to London by the roads of 1805 (Vallance et al., in prep.), 1t is appropriate to examine what befell the specimens after their arrival in London. They appear to have gone initially to Banks’s house in Soho Square. There and later at the British Museum, they were a major part of Brown’s herbarium (Murray, 1904). During his lifetime Brown kept his own herbarium separate from the British Museum material and did not allow free access (Edwards, 1976), but George Bentham [1800-1884] consulted it in the preparation of his Flora (Bentham and Mueller, 1863- 1878) (cf. Stearn, 1981). The next to feature as custodian of Brown’s plants was J. J. Bennett [1801-1876]. Bennett became assistant to Brown at the British Museum in November 1827 and Brown’s herbarium was left to him at Brown’s death (Woodward, 1904; Mabberley, 1985). In his turn Bennett kept the Brown herbarium separate from the main herbarium but on his death (February 1876) his wife gave it to the British Museum and it was then incorporated into the General Herbarium. Duplicates, and material which today would not necessarily be considered as duplicate, were later dis- tributed to other institutions in Europe and Australia. Those sent to Ireland have been listed by Powell and Morley (1976). Unlike the practice in other British Museum departments, the Botany Department never prepared formal registers, so there are no present-day BM (NH) herbarium numbers as such. There are indications that some Brown Herbarium vascular plant specimens were given to Banks’s librarian Jonas Dryander [1748-1810] during his lifetime. The Brown specimens listed in the Bennett register do not include these duplicates. In our list we have included them only where we believe the original is lost and where we can relate them to the Bennett register entries. These Dryander duplicates are recognizable by Dryander’s later label, but bear neither a Brown original label nor the blue sticker of Bennett’s plants (see below), nor a number. Some are unique while others are plants from the same gathering as that to which a Bennett number was given. The actual numbering was done by James Britten [1846-1924], assistant and later librarian, in the Botany Department and was probably undertaken soon after Brown’s herbarium was acquired on the death of Bennett. Not all species given by Brown (1810) are in our list, as Brown sometimes selected Banks and Solander specimens for his descriptions. As an example, Mitrasacme prolifera R.Br., Spigeliaceae (Bennett 2894) bears a Brown label indicating that it came from the Endeavour River, Queensland, in 1770. The description of this taxon in the Prodromus (Brown, 1810: 453, entry 8) then, is of a Banks and Solander gathering, not Brown's. Fortunately, Brown annotated his algal and pteridophyte original labels after the publication of Turner’s (1807-19) Fuci and his own Prodromus (Brown, 1810). So we have reason for thinking that certain specimens are illustrated, or at least noticed, in these works. A fern example is Bennett 104 (Gleichenia rupestris R.Br., see below). Bryophyte and gymnosperm labels are not annotated in this way. ‘THE BENNETT REGISTER After coming into possession of Brown’s Herbarium Bennett instituted a register of PROC. LINN. SOC. N.S.W., 111 (2), 1989 68 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS the material. This gave the plants both a number and a name. But it was left to Britten, assisted by Henry Trimen [1843-1896], to complete the task (Edwards 1976). No locality or ecological data are recorded in this ‘Bennett Register. Such detail as Brown com- mitted to paper is obtainable only from the original labels on the herbarium sheets, and, in some instances, from Brown’s descriptions in the Botany Library (the ‘Brown Slips’). In the herbarium the Brown specimens catalogued by Bennett and his followers are recognized by a blue label bearing Britten’s hand-written number and the printed words ‘Iter Australiense 1801-5”. One hundred and twenty-three numbers were allocated to Brown’s algae and charophytes in the Bennett register. Of these, we have been able to find eighty-four. Because Bennett’s workers missed specimens which we include, it is difficult to make numbers of ‘found’ taxa agree with the Bennett register. Some of Brown’s algae were described by Dawson Turner [1775-1858], and in the list below, dates of publication of parts of Turner's Fuci (1807-1819) are given according to Price (1984). Fortunately, the specimens described by Turner (1807-19) are recognizable because Brown annotated his labels to the effect ‘this 1s the specimen sent to Mr Turner’. The absence of Brown’s original labels in some cases means there is no locality information for some specimens. Also, the Brown locality ‘South Coast’ is disappoint- ingly vague. But better locality data are often available where a specimen is mentioned in Turner (1807-19). In our list nomenclature, where appropriate, is in accordance with Womersley (1967, 1984, 1987). The charophyte nomenclature used in our list is that of Wood (1965) and we have located four out of five of Brown’s charophytes. The lichens were never given space in the Bennett register. It would appear that they were handed in the 1870s to the Rev. J. M. Crombie [1830-1906] to investigate, probably with a batch of blank ‘Iter Australiense’ labels. It is Crombie’s writing on the blue labels accompanying Brown’s original lichen labels and it would appear that Crombie also applied the series of numbers. The numbers used are between 500 and 579 (thus not duplicating any numbers assigned by Bennett and Britten). It is these numbers which were quoted by Crombie (1879). It can be seen from the lichen list that there are several species given the same number. This is because since the 1870s many have received revision and so now consist of more than one species under one number. Many of Brown’s lichens were collected in Tasmania, and Kantvilas (1983) warns of the unreliable nature of many of Brown’s labels. But, as others have found (Vallance and Moore, 1982), the original labels may give reliable provenance data and should not be regarded lightly. Assuming that originally there were seventy-nine lichens, and each was allocated a number by Crombie, we have been unable to trace twenty-four of them. The fungi too are somewhat neglected by the Bennett register. They were allocated numbers 142-156 (inclusive), but no identifications or other data were entered against the numbers. The specimens, probably all mounted, were transferred to Kew Her- barium when the BM(NH) fungal collection was passed to Kew as a result of the Moreton agreement of 1961 (cf. Stearn, 1981: 309-310). They have since been incor- porated into the Kew collections. A search made there by one of us (EWG) to trace Brown’s fungi, with the aid of the list of the ten names in Brown’s appendix to Flinders (1814: 2, 592-4), has revealed only one possible fungus. This is ‘Clavaria coralloides L’ (= Clavaria cristata var. coralloides Corner). While not attributed to Brown on the sheet, the writing beneath the specimen and the watermark of the paper appear to be con- temporary with other Brown sheets at the BM(NH). The majority of Brown’s bryophytes in the BM(NH) remain unmounted and have still to be critically examined. As they lack Bennett numbers they have not been in- cluded here. But a minority were allocated Bennett numbers and subsequently PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 69 incorporated in the herbarium. These appear to be mostly from Tasmania. There is Brown material on some sheets from the Hooker herbarium (ex Kew) but this does not have original labels. From Brown’s Diary (cf. Vallance et al., in prep.) we know that Brown visited only Port Dalrymple, and the Esk Rivers and the River Derwent/Mt. Wellington areas of Tasmania in 1804. Despite this, we have accounted for all the spaces in the Bennett register assigned to bryophytes. In the 1870s Bennett assigned one hundred and forty-four numbers to Brown’s pteridophytes. We have found all but two. Brown (1810) indicated by letters the provenance of the species described, for example ‘D’ for Tasmania, and ‘]’ for Port Jackson. The original labels were annotated by Brown so there is reason for thinking that certain specimens are the ones that formed the basis of the descriptions in Brown (1810). Three cycadophytes are noted in the Bennett register. While we have traced two, there remains uncertainty over the identity of ‘Zamia spiralis’ of Flinders (1814: 1, 81). It appears to be a late entry to the Bennett register and thus existed in the 1870s. We have not traced it. Brown (1810: 348, entry 1 ‘J’ & ‘M’) indicated his specimen was not tropical and from study of Brown’s Diary (Vallance et a/., in prep.; cf. Flinders, 1814: 1, 348) we believe the specimen came from Lucky Bay, Western Australia. Consequently, we think it was, in fact, Macrozamia riedle: (Fisch. ex Gaudich.) C. A. Gardn., Zamiaceae (cf. Johnson, 1959: 103). This 1s ‘Macrozamia spiralis Miq’ of Mabberley (1985: 86). In our list of Brown’s conifers we follow the nomenclatural recommendation of Mr K. Hill of the Royal Botanic Gardens, Sydney. Bennett assigned nineteen spaces in his register to this group. We have found fourteen specimens. The manuscript Bennett register (now housed in the Botany Library, BM (NH) ), is the only document to list the plant specimens gathered by Brown in Australia. But it was written seventy years after the voyage ended and is a somewhat inadequate docu- ment for curatorial purposes. There were, for example, 3900 numbers allocated but con- siderably fewer than this number of actual specimens; there are blank numbers in the register and the numbers do not run consecutively. For example, the bryophytes finish at 295 and the next entry, the Asteraceae (Compositae), start at 2000. So, as the sequential arrangement in the Bennett register bears no chronological relation to the order of collection, and exhibits only a rudimentary grouping into families, there is no advan- tage in giving our list in this manner. Consequently in our list the genera are arranged in alphabetical order under their respective groups. The number we have given to each entry has been applied merely to aid reference. NOTES ON THE SPECIMEN LIST 1. In the following list we give Brown’s original label entry, with his geographical and ecological data as they are set out on the labels. Keys to the geographical localities and anchorages around Australia will be found in Burbidge (1956) and Stearn (1960a). 2. For some listed specimens there is more than one original label per herbarium sheet, but in numerous instances there is more than one sheet per label and more than one plant per sheet. In some cases it is not always clear which label goes with which plant. 3. We have kept editorial comment on the texts of Brown’s original labels (given here in single quotation marks) to a minimum, but we have indicated any references on the original labels to Turner’s Fucz (1807-1819), Crombie (1879) and Brown (1810). The note ‘Icones Bauer’ found on some higher-plant labels appears not to be present on the pteridophyte labels. Our claim that the Bauer painting of, for example, Azolla, is based on a particular herbarium specimen is circumstantial and visual. PROC. LINN. SOC. N.S.W., 111 (2), 1989 70 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS ALGAE CHLOROPHYTA 1. Anadyomene browni (Gray) J.Ag. Bennett 205 Lacks a Brown original label, and is thus unlocalized. An annotation on the sheet indicates that this is ‘Calomena brown1 of Gray (1866: t.44, fig. 3) and has type status. 2. Caulerpa browni (C.Ag.) Endl. Bennett 173 ‘Actaia selago? [sic] Kents Islands’ There are two sheets of this material with identical labels. 3. Caulerpa cactordes (Turn.) C.Ag. Bennett 166 ‘Actaia opuntia South Coast the specimen sent to Mr Turner’. ‘Fucus cactoides’ of Turner (1809-11: 3, 89-90, t.171). Type status. 4. Caulerpa flexilis Lam. Bennett 172 (Syn. ‘Caulerpa hypnoides’ (R.Br.) C.Ag.) ‘Actaia hypnoides Kents Islands’. Turner (1809-11: 3, 93-4, t.173). Two specimens, one with type status. 5. Caulerpa geminata Harv. Bennett 167 (Syn. ‘Caulerpa sedoides’ (R.Br.) C.Ag.) Two identical labels on the sheet. ‘Actaia sedoides L Kents Islands. The specimens sent to Mr Turner’ In fact, ‘Fucus sedoides’ of Turner (1809-11: 3, 92, t.172). There is an annotation on another sheet ex Turner herb. (formerly at Kew) by I. R. Price, 16 x11 1974 to the effect that the specimen was the basis of Turner’s figure. Type status. 6. Caulerpa racemosa var. laetevirens (Mont.) Weber van Bosse Bennett 168 No original labels, and specimen is consequently unlocalized, but a note by I. R. Price of 10 x11 1974 indicates he considered the material to be possibly of the tropical var. lamourouxit (Turn.) Weber van Bosse. 7. Caulerpa racemosa (Forsk.) J.Ag. var. laetevirens (Mont.) Weber van Bosse Bennett 169 ‘Actaia clavata South Coast’. I. R. Price in a note of 10 x11 1974 considers this locality incorrect. 8. Caulerpa racemosa (Forsk.) J.Ag. var. laetevirens (Mont.) forma cylindrica (Sonder) Weber van Bosse Unnumbered (Syn. ‘Caulerpa cylindracea’ Sonder) Two specimens on one sheet, the lower one is labelled ‘Actaia clavata South Coast’ 9. Caulerpa scalpelliformis (R.Br. ex Turn.) J.Ag. Bennett 171 Two labels: ist reads ‘Actaia scalpelliformis South Coast’; 2nd reads ‘Actaia scalpeliformis South Coast the spec sent to Mr Turner’ Turner (1809-11: 3, 95-6, t.174). Type status. 10. Caulerpa sertulariodes (S.Gmel.) Howe Bennett 170 (Syn. ‘Caulerpa taxifolia (Turn.) C.Ag.) Two sheets, label reads ‘Caulerpa taxifolia Fucus taxifolius Turn South Coast’. ‘Fucus taxifolius’ is featured in Turner (1807-8: 1, 120-122, t.54). In a note on the sheet of 10 xi1 1974 I. R. Price considered this taxon to be tropical. PROG. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 7 11. Caulerpa simpliciuscula (Turn.) C.Ag. Bennett 174 There are two sheets ‘Actaia conglobata Kents Islands. Turner 1809-11: 3, 98, t.175). 12. Codium australicum Silva Bennett 181 (pro parte) No Brown label, consequently unlocalized. 13. Codium duthieae Silva Bennett 181 (pro parte) No Brown label. 14. Codium fragile (Suringar) Hariot ssp. tasmanicum (J.Ag.) Silva Bennett 181 ‘SC’ [?South Coast]. (pro parte) 15. Codium mueller Kutz. Bennett 181a No Brown label and unlocalized. 16. Polyphysa peniculus (R.Br. ex Turn.) C.Ag. Bennett 203 (Syn. ‘Acetabularia peniculus’ (R.Br. ex Turn.) Solms-Laubach) Has type status but sheet on loan in November 1988. PHAEOPHYTA 17. Acrocarpia paniculata (Turn.) Aresch. Bennett 213 Two labels: Ist reads “The sp sent to Mr ‘Turner Fucus scoparino Kents Islands Bass Strait’; 2nd reads ‘Fucus paniculatus ‘Turn. hist 3 p.99 t.176 Cystoseira paniculata Agardh [1823-8] Sp. alg. 1 p.76. Turner (1809-11: 3, 99-100, t.176). Type status. 18. Asperococcus bulbosus Lam. Bennett 179 ‘F[ucus] ampullaceus is an incorrect name for a plant wch when perfect is filld with a dense gelatinous material’ [reverse reads] ‘Kents Islands & South Coast’. 19. Carpoglossum confluens (R.Br. ex Turner) Kutz. Bennett 226 ‘Fucus confluens Port Dalrymple’. ‘Fucus confluens’ of Turner (1809-1811: 3, 17, t.141), Agardh (1823-8: 1, 95). Type status. 20. Cystophora brown (Turn.) J.Ag. Bennett 208 ‘Fucus Browni Turn Hist 4 pl t 197 Cystoseira Brownii Agardh sp algar; 1 p.73’ [reverse] ‘sent to Mr Turner, you proposd to give it my name South coast frequent. According to Turner (1811-19: 4, 2-3), the specimen is from King George Sound. See also, Agardh (1823-8: 1, 73). H. B. Womersley appended a note, dated 31 v 1952, to the effect that the lower of the two specimens on the sheet is the type of ‘C. browni’, but the upper may be ‘C. monilifera’. 21. Cystophora moniliformis (Esper.) Womersley & Nizamuddin ex Womersley Bennett 214 Two labels, 1st reads ‘non logica F barbata sed distincta & prope F thuyoid: Herb Banks’ [reverse] ‘Fucus paniculatus litt. Kents Group’; 2nd reads ‘Fucus spartioides Turn hist. 4 p.85 t.232. Mr Turner did not return the specimen which is in fructification RB’ Turner (1811-19: 4, 86, t.232). Type status. 22. Cystophora retroflexa (Labill.) J.Ag. Bennett 209 Two labels: 1st reads ‘Fucus ramosissimus South Coast sent to Mr Turner’ 2nd reads PROC. LINN. SOC. N.S.W., 111 (2), 1989 72 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS ‘Fucus retroflexus Labill. Turn. hist. 3 p.47 t.155 Cistoseira retroflexa Agardh [1823-8] sp Alg. 1 p. 7#. Turner (1809-11: 3, 48, t.155). 23. Cystophora torulosa (R.Br. ex Turn.) J.Ag. Bennett 211 ‘Fucus torulosus Kents Islands Bass Strait 1803-4 The sp sent to Mr Turner. Turner (1809-11: 3, 51, t.157). Another sheet (ex Kew herbarium), lacks a Bennett number, but is almost certainly another portion of the same gathering. It has a label which reads ‘dup Fucus torulosus N Sp Turn fuc 3 [. . .] t157 Kents Group Cystophora torulosa Agardh’. Turner (1809-11: 3, 52, t.157). Type status. 24. Cystophora sp.indet. Bennett 219 ‘Fucus vagus the sp sent to Mr Turner Fucus vagus South Coast. The Turner illustration implied by the original label is untraced. 25. Cystophora sp.indet. Bennett 221 ‘Cystoseira (?) affin retroflexa Dr Agardh has a perfect specimen from Dr Greville. Unlocalized. 26. Cystophora sp.indet. Bennett 224 ‘Cystoseira |. . .] retroflexa. Unlocalized. 27. Cystosezra trinodis (Forsk.) C.Ag. Bennett 218 (Syn. ‘Cystophyllum muricatum’ (Turn.) J.Ag.) Two labels: Ist reads ‘Fucus angustifolius North Coast of New Holland F muricatus Mr Turner’s writing’; 2nd reads ‘Cystoseira muricata North Coast Cystoseira trinodis Ag [1823-8], Sp Alg p.67 [. . .] 24 June 1833’ Turner (1808-09: 2, 107-8, t.112). 28. Cystosezra trinodis (Forsk.) C.Ag. Bennett 223 No original labels, and unlocalized. 29. Dictyopterts woodwardia (R.Br. ex Turn.) J.Ag. Bennett 177 Two identical labels which read ‘Fucus woodwardia North Coast [reverse reads] a F membranaceo divers frondibus spinulo-denticula. A third sheet ex Herb. Kew, although lacking the Bennett number, is of the same gathering. Type status. 30. Dictyota sp.indet. Bennett 251 ‘Kents Islands’. 31. Ecklonia biruncinata (Bory) Papenfuss Bennett 229 (Syn. ‘Ecklonia radiata (Turn.) J.Ag.) ‘Fucus radiatus ‘Turn. hist 2 p.161 t.134 Port Dalrymple too young for Baptism [sic]. Turner (1808-09: 2, 161-2, t.134). An annotation on the sheet indicates this specimen was identified by H. B. S. Womersley 25. v. 1952 as the type of Turner’s var. exasperata of F. radiatus. 32. Homoeostrichus sinclairit (Hook. & Harv.) J.Ag. Bennett 175 (Syn. ‘Zonaria stuposa R.Br. ex J.Ag.) Two labelled sheets: 1st reads ‘Zonaria Novy sp’; 2nd reads ‘Kents Islands’. Type status. 33. Hormosira banksit (Turn.) Decaisne No Bennett number No sheet bearing Brown’s original label exists. There is, however, a label in Brown’s PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE the) hand ex Herb. Kew which reads ‘Nov. Holl. or occident. Herb. Mus. Paris Sep 1816 [added later] Cystoseira expansa Ag’ [a third sheet is labelled] ‘Cistoseira Banksii. Type status. 34. Macrocystus pyriforme (L.) C.Ag. Bennett 231 ‘Fucus pyriformis King’s Island’. 35. Pertthalia caudata (Labill.) Womersley Bennett 227 Two labels: 1st reads ‘Fucus inermis Port Dalrymple & South Coast of New Holland; 2nd reads ‘Fucus inermis distinct from F aculeatus with minute globules terminating the ramuli. Type status. 36. Phyllospora comosa (Labill.) C.Ag. Bennett 230 Two identical labels reading ‘Fucus polyphyllus South Coast of New Holland F comosus. We note ‘Turner (1809-11: 3, 18, t.142) gives the locality as ‘Van Diemens Land’. 37. Platythalia quercifolia (R.Br. ex Turn.) Sond. Bennett 217 Three labels: 1st reads ‘Fucus quercifolius South Coast New Holland 1802 [. . .]; 2nd reads ‘Fucus quercifolius South Coast of New Holland Turn hist 3 p37 t.151 Cystoseira quercifolia Ag’; 3rd label reads ‘Fucus quercifolius Turn hist 3. p.37 t.151. Turner (1809- 11: 3, 37, t.151). Type status. 38. Sargassum decipiens (Turn.) J.Ag. Unnumbered Two labels: Ist reads ‘Fucus polymorphus [struck through] decipiens Port Dalrymple the sp sent to Mr Turner’; 2nd reads ‘Fucus decipiens Ag Turn fuc 3 p79 t166. Turner (1809- bE 9> t166). Iivpe status: 39. Sargassum decurrens (Turn.) J.Ag. Bennett 157 ‘Fucus decurrens Turn hist 3 pl43 t194 North Coast (I believ) [sic] only specimen sp figured. See Turner (1809-11: 3, 143, t.194). Type status. 40. Sargassum heterophyllum C.Ag. Unnumbered Turner (1808-9: 2, 62, t.92) cites a Brown specimen of ‘Fucus heterophyllus’ from ‘Coast of New Holland’ but this material is untraced. 41. Sargassum lacerifolium (R.Br. Turn.) J.Ag. Bennett 158 ‘Fucus lacerifolius Turn hist 3 p81 t.167 Sargassum lacerifolium Agardh [1823- 8] sp alg. 1- p15 [. . .]. From Port Dalrymple, Tasmania, according to Turner (1809-11: 3, 81-2, t.167). Type status. 42. Sargassum paradoxum (R.Br.) Harv. Bennett 210 Two labels: Ist reads ‘Fucus paradoxus South Coast’; 2nd reads ‘Fucus paradoxus Mss Turn Fuc 3 p.49 t156 Cystoseira paradoxa Agardh [1823-8] in sp. alg. 1 p.15. Turner (1809-11: 3, 49, t.156). Type status. 43. Sargassum vestitum (R.Br. ex Turn.) C.Ag. Bennett 159 Two labels: 1st reads ‘Fucus vestitus Kents Islands Bass strait & South Coast The speci- men sent to Mr Turner; 2nd reads ‘Fucus vestitus Turn. hist. 3 pl01 t177 Sargassum vestitum Agardh [1823-8] Sp. Alg. 1 p24. Turner (1809-11: 3, 101-2, t.177). Type status. PROG. LINN. SOG. N.S.W., 111 (2), 1989 74 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 44, Sargassum sp.indet. Bennett 161 ‘Sargassum North Coast of New Holland’. 45. Sargassum sp.indet. Bennett 162 ‘Sargassum cum S. ilicifolia receptaculis convenit North Coast’ 46. Sargassum sp.indet. Bennett 163 ‘Sargassum North Coast of New Hollnad’ 47. Scytothalia dorycarpa (Turn.) Grev. Bennett 216 Two sheets and two labels: 1st reads ‘Fucus platycarpus South Coast of New Holland The specimen sent to Mr Turner; 2nd reads ‘Fucus dorycarpus Turn hist 3. p.21 t.143 Cystoseira dorycapa Agardh [1823-8] Sp alg 1 p.80°. ‘It is not clear in Turner (1809-11: 3, 24, t.144) ifthe illustrated specimen was collected by Menzies or Brown. 48. Seirococcus axillaris (R.Br. ex Turn.) Grey. Bennett 215 Two sheets, three labels in all: 1st reads ‘Fucus axillaris Port Dalrymple The sp sent to Mr Turner sp fig’; 2nd reads ‘Fucus axillaris Turn hist 3 p.27 tl46 Cystoseira axillaris Agardh [1823-8] sp. alg p.80’; 3rd reads “Cystoseira axillaris Port Dalrymple’. ‘Turner (1809-11: 3, 28, t.146). Type status. 49. Splachnidium rugosum (L.) Grev. Bennett 225 Two labels: 1st reads ‘Fucus rugosus Kents Islands shallow water on rocks. You say [sic] Ulva rugosus Herb/?] Lad’; 2nd reads ‘Fucus rugosus Turn. hist 3 p.119 t.185 Agardh [1823-8] Sp alg. 1 pl100 Kents Islands’ Turner (1809-11: 3, 118, t.185). 50. Sporochnus radiciformis (R.Br. ex Turn.) C.Ag. Bennett 228 ‘Fucus ?radiciformis is ita Fucus Kents Islands Bass Strait. Type status. 51. Xiphophora chondrophylla (R.Br. ex Turn.) Montague ex Harv. Bennett 271 Two labels: ist reads ‘Fucus chondrophyllus Port Dalrymple’; 2nd reads ‘Fucus chondrophyllus Mss Turn. hist 4 p. 60 t.222” Turner (1811-19: 4, 60, t.222). 52. Zonaria sp.indet. Bennett 182 No Brown label and consequently unlocalized. RHODOPHYTA 53. Brongniartella australis (J.Ag.) Schmitz Bennett 195 ‘Griffithsia australis Ag [. . .]. Unlocalized. Agardh (1823-8: 2, 135). 54. Champia parvula (C.Ag.) Harv. Bennett 240 No orig. label, consequently unlocalized. 55. Coelarthrum muellert (Sonder) Borges. Bennett 245 ‘Fucus compar Kents Islands Bass Strait & South Coast of New Holland’. Another sheet ex Herb. Kew, of the same gathering, bears the locality in Brown’s handwriting as ‘South & East coasts of New Holland’. PROG. LINN. SOG. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 75 56. Gelidiella acerosa (Forsk.) J. Feldmann & Hamel Bennett 264 (Syn. ‘Gelidiopsis rigida’ Grev. ) ‘Spharococcus rigidus Ag [1823-8] Sp. Alg. p.285 [. . .]. No locality or other data on original label. 57. Gelidium australe J.Ag. Bennett 250 ‘F corneus? [sic]. No locality given. 58. Gelidium glandulaefolium Hook. & Harv. Bennett 272 No Brown label. Secondary label reads ‘Fucus Kents Group’. 59. Gracilaria edulis (S.Gmel.) Silva Bennett 269 (Syn. ‘Gracilaria corniculata (R.Br. ex Turn.) J.Ag.) Two labels: Ist reads ‘Fucus ceratophyllus [struck through] corniculatus The sp sent to Mr Turner Kents Islands 1803-4 proxe cornea a distinc: spec? [sic]; 2nd reads ‘Fucus corniculatus Turn 3 p.113 t.182. Turner (1809-11: 3, 112, t.182). A note appended by H. B. S. Womersley on 27.v.1952 indicates ‘this specimen may well be a Gigartina. ‘Type status. 60. Griffithsia sp. indet. Bennett 193 ‘Griffithsia. No locality given. 61. Hymenocladia divaricata (R.Br.) Harv. Bennett 254 ‘Fucus divaricatus Mss in fructific only specimen Fucus divaricatus Turn hist 3 p.110 t181. Turner (1809-11: 3, 110, t.181) localizes Brown’s material as from ‘on the coast of New Holland’ ‘Type status. 62. Hymenocladia usnea (R.Br.) J.Ag. Bennett 259 ‘Fucus usnea Mss Kents Group Basses Strait 1803-4. A second label, not Brown’s, reads ‘Fucus usnea Ag Turner Hist 4 p70 t225. Chondria usnea Agardh [1823-8] Syn Alg. XVIIT. Turner (1811-19: 4, 70, t.225). Type status. 63. Hypnea musciformis (Wulf.) Lamour. Bennett 236 ‘F musciformis. Unlocalized. 64. Laurencia botryowdes (Turn.) Gaill. Bennett 253 On loan at time of compilation. 65. Laurencia pinnatifida (Huds.) Lam. Bennett 257 Two sheets with identical labels; ‘Fucus pinnatifidus Kents Islands’. 66. Leptophyllis conferta (R.Br. ex Turn.) J.Ag. Bennett 263 Two labels: ist reads ‘Fucus confertus Kents Islands 1803 Scopula nov sp? [sic]’; 2nd reads ‘Fucus confertus Mss Turn. 3 p177 t184 Delesseria ?conferta Ag. [1823-8] Sp. alg. p. 177? [sic]. Turner (1809-11: 3, 116, t.184). Type status. 67. Melanthalia obtusata (Labill.) J.Ag. Bennett 260 Two labels: ist reads ‘Fucus strictus Port Dalrymple F obtusatus Labill’; 2nd reads ‘Fucus obtusatus Labill Turn. hist 3 p.25 t.145. Turner (1809-11: 3, 25-6, t.145). PROG. LINN. SOG. N.S.W., 111 (2), 1989 76 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 68. Osmundaria prolifera Lamour. Bennett 206 (Syn. ‘Polyphacum proliferuny C.Ag.) ‘Polyphacum proliferum Agardh [1823-8] Sp. alg. 1. p.106. Osmundaria prolifera Lamour Port Dalrymple the only specimen’ 69. FPeyssonnelia australis Sond. Bennett 176 No Brown label, consequently unlocalized. 70. Phacelocarpus labillardiert (Mert. ex Turn.) J.Ag. Unnumbered No Brown gathering of this taxon has been traced in the BM (NH) herbarium, but there is a specimen marked ‘Australia R Brown’ (not in Brown’s writing) and a fragment on another sheet labelled ‘F scolopendrum Mss Mr Brown 1807 [sic]. Both sheets were formerly in the Kew herbarium. ‘Turner states regarding ‘Fucus labillardierr that “This plant has also been gathered upon the coast of New Holland by our own countryman, Mr Brown’ Turner (1809-11: 3, 8, t.137). 71. Plocamium sp.indet. Bennett 251 ‘Fuci pulchilli var? [y] Junior planta? Kents Islands Bass Strait according to you may be F aspenoides in herb’ Secondary label reads “Delessaria plocamum [a]. 72. Plocamium sp.indet. Bennett 252 ‘Fucus pulchellus Kents Island in Bass Strait & Port Dalrymple 1803-4. 73. Polyopes constrictus (Turn.) J.Ag. Bennett 268 ‘Two labels: Ist reads ‘Fucus densus Kents Islands 1803-4’; 2nd reads ‘Fucus constrictus Ain: hist 3 p39 tla27 See lurmer (1809-115 3) 39=40) tlb2)) type status: 74. Folysiphonia decipiens Mont. Bennett 235 (Syn. ‘Polysiphonia cancellata Harv.) Lacks original label, and consequently unlocalized. 75. Pterocladia lucida (R.Br. ex ‘lurn.) J.Ag. Bennett 266 ‘Fucus lucidus Mss ‘Turn. hist. 4 p99 t238. ‘Turner (1811-19: 4, 99, t.238) gives the locality as ‘on the southern coasts of New Holland’. 76. Rhabdonia verticillata Herv. Bennett 244 ‘Fucus 1s it difft from Compar [sic] Kents Islands’. 77. Rhodymenia cunerfolia (Hook.f. & Harv.) Taylor Bennett 249 (Syn. ‘R. corallina’ Grey.) No Brown label, consequently unlocalized. 78. Trematocarpus concinnus (R.Br.) De ‘Toni Bennett 233 ‘Fucus caespitosus according to you a var. of F plicatus Kents Islands One spec in fructif. ‘Type status. 79. Trematocarpus concinnus (R.Br.) De Toni Bennett 267 Two labels: 1st reads ‘Fucus scoparius [struck through] concinnus Kents Islands 1803-4; 2nd reads ‘Fucus concinnus Mss ‘turn. hist 3 p41 t153. See Turner (1809-11: 3, 41-2, t.153). Type status. PROG. LINN. SOG. N.S.W,, 111 (2), 1989 E. W. GROVES AND D. T. MOORE 77 80. Vidalia fimbriata (R.Br.) J.Ag. Bennett 261 Two labels: Ist reads ‘Fucus fimbriatus North Coast of New Holl spec unic (?){sic]’; 2nd reads ‘Fucus fimbriatus Mss ‘Turn. Hist. 3. p87 t170’. Turner (1809-11: 3, 87-8, t.170). Type status. CHAROPHYTA 81. Chara corallina Klein ex Willd. Bennett 277 (Syn. ‘Chara australis’ R.Br.) ‘Chara crassifolia Port Jackson’ 82. Chara corallina Klein ex Willd. Bennett 279 (Syn. ‘Chara plebeja R.Br.) ‘Chara A Carpentaria Point [sic, locality not given]. 83. Nitella congesta(R.Br.) A.Br. Bennett 276 ‘Chara cfr B a Bay I South Coast’. Lucky Bay, Western Austalia (cf. Burbidge, 1956). 84. Nitella hyalina (DC.) J.Ag. Unnumbered Two sheets with identical data ‘Chara B Point T Carpentaria. Cape Shield, Northern Territory (cf. Burbidge, 1956). LICHENES 85. Cladia aggregata (Sw.) Nyl. Bennett 532 ‘Lichen multiflorius Cladonia multiflora ericet elevat aust:’ (cf. Crombie, 1879: 393), a type according to Filson (1981: 15). 86. Cladia ferdinand (Mull. Arg) R. Filson Bennett 533 ‘Lucky Bay Cladia retipora. Another sheet without a Bennett number, but of the same gathering, bears the label ‘On the ground Lucky Bay Tasmania [sic]. A note by R. Filson of 8 vi 1983 states that ‘this specimen was probably collected at Lucky Bay, W Australia [...]’. The identification of this material has been confirmed by D. J. Galloway as Cladia ferdinandit. According to Crombie (1879: 393) Brown’s ‘Lichen cribrosus’ was from ‘Mountain heaths, Australia’. 87. Cladta retipora (Labill.) Nyl. Bennett 534 ‘Lichen cribrosus Cladonia cribrosa enceta Montes que ora australis & Ins Diemen’ Tasmania, cf. Crombie (1879: 393) and Filson (1981: 23). 88. Cladia retipora (Labill.) Nyl. Bennett 534 No original label, probably another part of the gathering above. 89. Cladonia squamosa Mill. Arg. Bennett 531 (Syn. ‘Cladonia subsquamosa’ (Nyl.) Wain var. ‘pulverulenta (R.Br.) Vain. and ‘C. pulverulenta R.Br.) No original label; Crombie’s handwriting reads ‘on the ground Table Mt. Tasmania R. Brown Cladonia acuminata (Achy. Brown’s ‘Cladonia pulverulenta was from Mt Wellington (cf. Crombie 1879: 392). PROC. LINN. SOC. N.S.W., 111 (2), 1989 78 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 90. Cladonia pleurota (Florke) Schaerer Bennett 530 (Syn. ‘Cladonia deformis’ Hoffm.) ‘lat. sax. Mont. Tabul prom Dieman’ Brown’s ‘Lichen deformis’ was from Mt Welling- ton, Tasmania, cf. Crombie (1879: 393). 91. Cladina confusa (R. Sant.) Follm. & Ahti Bennett 527 ‘Lichen rangiferinus mons Tabul & encet’.. Crombie (1879: 393) gives ‘Mt Wellington on the ground. 92. Coenogonium implexum Nyl. Bennett 504 ‘Lichen spongiosus arb cort: laevi in sylvis umbrosus humidis ad radis Mont Tabul prom Diemen’. Mt Wellington, Tasmania (cf. Crombie, 1879: 399). 93. Collema durvetzu Degel. Bennett 511 ‘Lichen fascicularis? rip & arb Musc insula Diemen’. This specimen had been earlier identified by Crombie as ‘Collema implicatum’ Nyl. From trees near the River Derwent, ‘Tasmania according to Crombie (1879: 391). 94. Collema leucocarpum Hook.f. & Tayl. Bennett 549 (bis) ‘Lichen obductus rup & arb orce australis N Holl & Ind Dieman’. Crombie (1879: 391-2). See also entry below. 95. Collema subconveniens Nyl. Bennett 549 (bis) We have not been able to trace this specimen, but Degelius (1974: 139) appears to have seen and cited it ‘as C. nigr[esens] Lichen obductus (BM) N.W. Tasm’ According to Crombie (1879: 392) Brown’s ‘Lichen obductus’ was from trees near the River Derwent, ‘Tasmania. 96. Diuploschistes ocellatus (Vill.) Norman Bennett 563 ‘Lichen lateritus [. . .] prope Risdon Cove’. Tasmania (cf. Crombie, 1879: 399). 97. Dirinaria picta (Sw.) Clem. & Shear Bennett 546 No original label; secondary label reads ‘Lichen pictus Sw near Kingstown, Newcastle’ (cf. Crombie, 1879: 397). 98. Ephebe fruticosa Henssen Bennett 522 Two species on sheet: label reads ‘Lichen pubescens Lin Ach L. exilis Lightf rup & sax inundat’. Specimen from the Grose River, NSW, according to Crombie (1879: 391). 99. Ephebe tasmanica Cromb. Unnumbered According to notes on the sheet there are three species present. The first is Ephebe tas- manica Cromb. The second Ephebe fruticosa A. Henssen. The third is Pseudephebe pubescens (L.) M. Choisy. (Syn. ‘Electoria pubescens(L.) Howe). According to Crombie (1879: 391) ‘Lichen pubescens’ was from River Grose, NSW. 100. Flavoparmelia rutidota (Hook.f. & Tayl.) Hale Bennett 539 ‘Lichen cfr L caperatum fl Derwent. Tasmania (cf. Crombie, 1879: 394). 101. Graphis intricata Fée Bennett 561 ‘Lichen Opegrapha dendritica arb: trunce rip: fl Grose [. . .]. According to Crombie PROG. LINN. SOG. N.S.W,, 111 (2), 1989 E. W. GROVES AND D. T. MOORE 79 (1879: 401), the specimen was from the bark of trees, near the River Grose, New South Wales. 102. Graphis platycarpa (Eschw.) A. Zahlbr. Bennett 560 ‘Lichen Opegrapha plebja cor: arb: prope P Jackson. (Graphis sophistica Nyl. and Opegrapha plebja of Crombie, 1879: 401). 103. Haematomma puniceum (Ach.) Mass. var. collatum (Stirt.) Zahlbr. Bennett 569 (Syn. ‘Haematomma collatum (Stirt.) Dodge) ‘Ad cortices Port Jackson. Probably “Lichen guttatus’ gathered from bark of trees near Middle Harbour, Port Jackson; cf. Crombie (1879: 399). 104. Haematomma puniceum (Ach.) Mass. var. infuscum (Stirt. ex Bailey) Zahlbr. (Syn. ‘Haematomma infusum (Stirton ex Bailey) R. W. Rogers) Bennett 507 ‘Lichen guttatus Middle Harbour Port Jacksor (cf. Crombie, 1879: 399). 105. Hypogymnia lugubris (Pers.) Krog Bennett 526 ‘Lichen nem aff L physodi lat Mont Tabularis’. Mt Wellington, Tasmania (cf. Crombie, 187925395): 106. Hypogymnia mundata (Nyl.) Rassad. Bennett 550 (bis) According to an annotation by J. A. Elix (of x 1977) this specimen, noted by Crombie as ‘Parmelia mundatum var. pulverata Nyl. [. . .] Lichen dendrosme on trees base of Table Mt, Tasmania, is a mixture of the above species and H. pulverata (Nyl. ex Crombie) Elix. 107. HAypogymnia pulverata (Nyl. ex Cromb.) Elix Bennett 515 (on some sheets as 550) No original label, but has been written up by Crombie (1879) as a mixture of Parmelia mundata Nyl. (ex Derwent River) and “Parmelia angustata’ Pers. (ex Port Jackson). Remounting seems to have removed P mundata, and the latter has been identified by J. A. Elix (x 1977) as the above. 108. HAypogymnia tabularis (Taylor) Elix Bennett 550 ‘Lichen dendrosma [...] in sylvis umbrosis ad radic Montis Tabularis prope fl Derwent. Determined by Crombie (1879: 395) as ‘Parmelia mundata Nyl., more recently by J. A. Elix (ix 1977) as the above. 109. Lecidea flindersii Cromb. Bennett 565 ‘Superce [?] terra Risdon Cove’. Tasmania (cf. Crombie, 1879: 400). 110. Melaspilea corcumserpens Nyl. Bennett 566 ‘Lichen ambiguens I terra nude rarius in saxis In collibus prope Risdon Cove’ Tasmania (cf. Crombie, 1879: 401). 111. Nephroma cellulosum (Ach.) Ach. Bennett 543 ‘Lichen antarcticus rup umbrosos humid adrip fl Anna Maria infra calaratumnY ‘Anna Maria River appears to be Margate Rivulet, southeastern Tasmania, (cf. Vallance et al., in prep.). 112. Pannaria rubiginosa (Thunb. ex Ach.) Del. Bennett 556 ‘South coast Bay 3”. Fowlers Bay, South Australia (perhaps Crombie, 1879: 397). PROC. LINN. SOG. N.S.W., 111 (2), 1989 80 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 113. Parmelia tenuirima Hook.f. & Tayl. Bennett 537 Two sheets: Ist reads ‘Lichen amplissimus in saxis & rupibus collium prope fl: Derwent’ 2nd sheet has no orig. label. “Lichen amplissimus’ of Crombie (1879: 394). 114. Pseudephebe pubescens (L.) M. Choisy Bennett 521 ‘Lichen lanatus sum Mont Tabul prom Diemen (cf. Crombie, 1879: 395). 115. Pseudocyphellaria billardiere: (Delise) Rasanen Bennett 536 ‘Lichen linearis in trunc arbor prope fluv Derwent Tasmania (cf. Crombie, 1879: 396). 116. Pseudocyphellaria crocata (L.) Vainio Bennett 540 (bis) ‘Lichen crocatus Dicks: Hoffm: rup & Sax in montosis’ (cf. Crombie, 1879: 396). See P neglecta, below, for second label. 117. Pseudocyphellaria glabra (Hook.f. & Tayl.) Dodge Bennett 538 ‘Lichen latissimus rup & sax Montis Tabularis. Mount Wellington, Tasmania (cf. Crombie, 1879: 396). Determined as ‘Sticta freycinetii’ Del. by Crombie. 118. Pseudocyphellaria multifida (Nyl.) D. Galloway & P. James Bennett 516 ‘Lichen dichotomus rup: umbros: |. . .] ad rep fl Anna Maria prope Calaractam (cf. Crombie, 1879: 396). Perhaps Margate Rivulet, Tasmania. 119. Pseudocyphellaria neglecta (Mull. Arg.) H. Magn. Bennett 540 (bis) ‘Lichen crocatus. Locality as 116 above? 120. Psoroma decipiens (Hedw.) Hoffm. Bennett 567 (2Syn. ‘Lecidea decipiens’) ‘Lichen decipiens [. . .] supra terra Risdon Cove’ Tasmania (cf. Crombie, 1879: 400). 121. Psoroma asperellum Ny)l. Bennett 547 ‘Lichen adscendens [. . .] ad latus Mont: Montis Tabularis. Mt Wellington, Tasmania (cf. Crombie, 1879: 398). 122. Psoroma hypnorum (Vahl) S. F. Gray Bennett 354 ‘Lichen cfr: L hypnorum lat sax: Montis Tabularis prope Derwent’. Mount Wellington, Tasmania. 123. Psoroma leprolomum (Nyl.) Rasanen Bennett 553 No original label, secondary label suggests specimen is ‘Lichen denrosmae’ from base of Mt Wellington, Tasmania, cf. Crombie (1879: 398). 124. Psoroma soccatum R.Br. ex Crombie Bennett 502 ‘Lichen soccatus trunce emiort ad radi Mont Tabularis’ (cf. Crombie, 1879: 398-9). Lectotype (Galloway, 1985: 481). 125. Psoroma sphinctrinum (Mont.) Nyl. Bennett 553 ‘Lichen Dendrosma reniles Derwent River’ Tasmania (?Crombie, 1879: 398). 126. Ramalina inflata (Hook.f. & Tayl.) Hook.f. & Tayl. Bennett 519 ‘Lichen fastigiatus arb [. . .] prope P Jacksor’ (cf. Crombie, 1879: 393). PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 81 127. Ramalodium succulentum Nyl. ex Cromb. Bennett 551 ‘Lichen succulentus arb: trunc & rup umbros ad rip fl Grose [. . .] 2 rami and musci (cf. Crombie, 1879: 392). 128. Relicina limbata (Laurer) Hale Bennett 544 Two specimens, but one original label missing; secondary label suggests ‘Lichen pin- natus on rocks from Port Jackson. 2nd (original) labels reads ‘Lichen pinnatus in saxis prope P Jackson’ (cf. Crombie, 1879: 394). 129. Siphula coriacea Nyl. Bennett 501 ‘Lucky Bay S Ct. Another sheet, without a Bennett label, bears a label in Crombie’s hand ‘on the ground Lucky Bay ‘Tasmania [sic]. See also number 86, above. Crombie (1879: 393), however, has number 501 from Mt Wellington, Tasmania. 130. Sphaerophorus melanocarpus (Sw.) DC. Bennett 517 Two sheets: 1st label reads ‘Lichen fragilis arb trunc semi putrid ad rip flu alpinor Mon Tabul prom Diemen’?; 2nd has no original label (cf. Crombie, 1879: 392). 131. Sphaerophorus tener Laurer Bennett 524 ‘Lichen globiferus lat sax Mont Tabul prom Diemen (cf. Crombie, 1879: 392). 132. Stereocaulon ramulosum (Sw.) Rauschel Bennett 526 ‘Lichen ramulosus men alt L paschali lat Mont Tabul &c rip fl Grose &c’ (cf. Crombie, 1879: 392). 133. Stricta subcaperata (Nyl.) Nyl. Bennett 555 ‘Lichen filix Lin: fil. In lat umbros supra summitato Montis Tabularis insul. Dieman’. Mount Wellington, ‘Tasmania (cf. Crombie, 1879: 396). 134. ‘Teloschistes chrysophthalmus (L.) Th. Fr. Bennett 545 ‘Lichen chrysophtphalamus [. . .] Kents Group.’ Bass Strait (cf. Crombie (1879: 396-7). 135. Thamnolia vermicularis (Sw.) Ach. ex Schaerer Bennett 528 ‘Lichen vermicularis Cladonia vermicularis lat & sum: Mon Tabul prom Diemen. Mount Wellington, Tasmania (cf. Crombie, 1879: 393). 136. Thelotrema lepadinum (Ach.) Ach. Bennett 520 ‘Lichen ? [sic] occulatatus Cort laevi: arbor ad ripas fluv: Grose [. . .]’ (cf. Crombie, 1879: 399). 137. Unmbilicaria cylindrica (L.) Delise in Duby Bennett 558 Two specimens: Ist has no original label, secondary label suggests ‘Lichen proboscideus summit of Table Mtn? 2nd (original) reads ‘Lichen proboscideus [.. .] sum: Mont Tabularis prope fl Derwent. Mount Wellington, Tasmania (cf. Crombie, 1879: 397). 138. Usnea acromelana Stirt. var. decipiens (Lamb) Lamb Bennett 523 (?Syn. ‘Neuropogon acromelanus’) ‘Lichen ustulatus rup sum: Mont Tabularis prom Diemen’ (cf. Crombie, 1879: 395). Holotype of variety (Walker, 1985). PROG. LINN. SOG. N.S.W., 111 (2), 1989 82 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 139. Usnea ceratina Ach. Bennett 518 (bis) (cf. Crombie, 1879: 394). Specimen untraced. 140. Usnea florida (L.) Wigg. Bennett 518 (bis) ‘Lichen floridus’ from the River Grose, N.S.W., specimen untraced. (cf. Crombie, 1879: 393). 141. Xanthoparmelia australiensis (Cromb.) Hale Bennett 525 ‘Scyphophorus lat Mont Tabul In Diem [sic]’. Mt Wellington, Tasmania according to Crombie (1879: 395), but this locality could be an error (Kantvilas 1983). According to Bibby (1951), and Ehix et al. (1986) the locality is the Nullarbor region and Flinders Ranges. Holotype of species (Elix et al., 1986). 142. Xanthoparmelia multipartita (Cromb.) Hale Bennett 542 ‘Lichen multipartitus in saxis prope P Jackson’. (cf. Crombie, 1879: 394-5). There is other material in the gathering with no original label; secondary label reads ‘Lichen multipartitus rocks near Port Jackson’ Taxon distributed in south-eastern Australia and this number is described as type by Elix et al. (1986). BRYOPHYTA LEAFY LIVERWORTS 143. Jamesoniella colorata (Lehm.) Spruce ex Schiffn. Bennett 293 Secondary label reads “Iasmania. ‘The taxon 1s known to occur in Tasmania according to Scott (1985). 144. Frullania falciloba Yaylor ex Lehm. Bennett 290 Secondary label reads “Tasmania. Known in Tasmania (Scott, 1985). 145. Gackstroema weindorfi (Herzog) Grolle Bennett 294 Secondary label reads “Tasmania. Known to occur, frequently in Nothofagus forests, in Tasmania (Scott, 1985). 146. Lepicolea scolopendra (Hook.) Dum. ex Trev. Bennett 292 Secondary label reads “Tasmania. 147. Plagiochila sp.indet. Bennett 291 Secondary label reads “Tasmania. ‘THALLOID LIVERWORTS 148. Hymenophyton flabellatum (Labill.) Dum. ex Trev. Bennett 295 (Syn. ‘Symphyogyna flabellata’ (Labill.) Mont.) Secondary label reads “Iasmania. Known in ‘Iasmania, Scott (1985). The specimen is unmounted, and in a folded and re-used sheet of Brown’s herbarium drying paper on which is written ‘Carpentaria Island s[. . .] 21 Jany 1803 [. . . but this is struck through in pencil. PROG. LINN. SOG. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 83 149. Symphyogyna podophylla (Thunb.) Mont. & Nees Bennett 296 (Syn. ‘Symphyogyna obovata Hook.f. & Tayl.) Secondary label reads “Tasmania. The label is annotated ‘Syn Hep p.480’ (cf. Gottsche et al., 1844-7). Scott (1985) confirms the Tasmanian occurrence of this taxon. MUSCI 150. Blindia magellanica Schimp. Bennett 284 Secondary label reads “Tasmania. Tasmanian occurrence is confirmed by Scott and Stone (1976). 151. Breutelia affinis (Hook.) Mitt. Bennett 288 Secondary label reads “Tasmania (cf. Scott and Stone, 1976). 152. Breutelia pendula (Sm.) Mitt. Bennett 289 Secondary label reads “Tasmania (cf. Scott and Stone, 1976). 153. Bryum billardiert Schwaegr. Bennett 287 Secondary label reads “Tasmania (cf. Scott and Stone, 1976). 154. Campylopus clavatus (R.Br.) Wils. Bennett 282 Secondary label reads “Tasmania (cf. Scott and Stone, 1976). 155. Campylopus introflexus (Hedw.) Brid. Bennett 281 Secondary label reads “Tasmania (cf. Scott and Stone, 1976). 156. Dawsonia polytrichoides R.Br. Unnumbered Ist original labels reads: ‘Nova Cambria Australis: ad radices montium prope Port Jackson Jan 1 1804 RB [sic]’; 2nd original label reads ‘Dawsonia polytrichoides to show that part of the peristomium originates from the columnale or Placenta. But an anno- tation on the wrapping paper of an unmounted specimen in a ‘Herb. Brown’ cover reads ‘Genus Muscor frondos Proxim: Polytricho lesser division of the first branch of the River Grose Jan 1 1805 desc’. The 1st original label is incorrectly dated. Brown was in Tasmania 1 January 1804 (cf. Vallance et al., in prep.). The date 1 Jan 1805 is consistent with the New South Wales locality. Probably material from this gathering was described by Brown (1811) where the locality was given as ‘Port Jackson. Scott and Stone (1976) note the occurrence of this taxon in Tasmania, Victoria, New South Wales, A-C:T. and Queensland. 157. Dhicranoloma dicarpum (Nees) Par. Bennett 285 Secondary label reads “Tasmania (cf. Stone and Scott, 1976). 158. ‘Dicranoloma robustum’ of BM (NH) Herbarium Bennett 283 Secondary label reads “Iasmania’. 159. Leptostomum erectum R.Br. Unnumbered Secondary label reads ‘New Holland Mr Brown’. The mounted Hooker specimen (ex Kew Herbarium) bears a secondary label which reads ‘New Holland Mr Brown’. Taxon known in Victoria, New South Wales and Queensland (Scott and Stone, 1976). Possibly gathered with Brown’s (1811) material from the ‘Hawkesbury and Grose’. PROG. LINN. SOG. N.S.W., 111 (2), 1989 84 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 160. Leptostomum inclinans R.Br. Bennett 286 Possibly this material was gathered with that described by Brown (1811) and from ‘Van Diemans Land’ A Hooker specimen of this gathering (ex Kew Herbarium) has a second- ary label which reads “Table Mountain Van Diemens Land Mr Brown at an elevation of 3000-3500ft. A label on the sheet indicates this specimen 1s the basis of the illustration by Brown (1811: 320, t.23, fig 2). A secondary label with the unmounted material reads “Tasmania (cf. Scott and Stone, 1976). 161. Leptostomum macrocarpum (Hedw.) Pyl. Unnumbered Secondary label reads ‘Leptostomum macrocarpum, Br Australia R Brown. Known from Tasmania, New South Wales and Queensland according to Scott and Stone (1976). 162. ‘ayloria octoblepharum (Hook.) Mitt. Bennett 280 Secondary label reads “Tasmania. Scott and Stone (1976) confirm the occurrence in Tasmania. PTERIDOPHYTA LYCOPSIDA 163. Lycopodium deuterodensum Herter Bennett 126 (on same sheet as Bennett 127) ‘Lycopodium candelabrum Derwent Huon &c (Lycopodium densum of Brown, 1810: 165, entry 3-D). 164. Lycopodium deuterodensum Herter Bennett 127 (on same sheet as Bennett 126) ‘Lycopodium venustrum Port Dalrymple Jan 1804 [reverse] Lycopodium candelabrum’. 165. Lycopodium deuterodensum Herter Bennett 128 ‘Lycopodium candelabrum B & S Port Jackson [reverse] 3 Lycopodium densum’ (Brown, 1810: 165, entry 3-J). 166. Lycopodium fastigiatum R.Br. Bennett 129 Two labels, ist reads ‘4 Lycopodium fastigiatum Summit of Table Mountain VDL (Brown, 1810: 165, entry 4-D); 2nd reads ‘Lycopodium quadrifarium Port Jackson’. 167. Lycopodium laterale R.Br. Bennett 130 ‘Lycopodium diffusum Port del Esperance V D Land’ (Brown, 1810: 165, entry 5-D). 168. Lycopodium laterale R.Br. Bennett 131 ‘Lyopodium laterale [. . .] Port Jackson’ (Brown, 1810: 165, entry 6-J). 169. Lycopodium myrtifolium Forst.f. Bennett 123 ‘Lycopodium varium [alpha]|[. . .] Table Mountain’ (Brown, 1810: 165, entry 1-D). 170. Lycopodium myrtifolium Forst.f. Bennett 124 ‘Lycopodium varium [beta] alpinae Mont Tabule’ (Brown, 1810: 165, entry 1-D). Holo- type of ‘Lycopodium varium [beta] alpinum’, according to an annotation on the sheet by J. M. Beitel 1981. 171. Lycopodium scariosum Forst.f. Bennett 125 ‘Lycopodium decurrens affr L reptans B & S: Mont Tabularis and fluy Derwent’ (‘Lycopodium decurrens’ of Brown, 1810: 165, entry 2-D). PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 85 172. Selaginella ulrginosa (Labill.) Spring Bennett 133 ‘Lycopodium quadrilineum var Port Phillip’ 173. Selaginella uliginosa (Labill.) Spring Bennett 132 ‘Lycopodium uliginosum Labill’ (Port Jackson, cf. Brown, 1810: 165, entry 7-J). PSILOPSIDA 174. Psilotum nudum (L.) Beauv. Bennett 120 Two labels: 1st reads ‘Lycopodium nudum Broad Sound [reverse] 1 Psilotum triquetrum’; 2nd reads ‘Lycopodioides nuda P.Jacksor (‘Psilotum triquetrum’ of Brown, 1810: 164, entry 1-J). 175. Timesipteris billardiert Endl. Bennett 122 (on same sheet as Bennett 121) ‘Lycopodioides truncata [. . .]| Derwent [reverse] 2 Psilotum truncatum’ (Brown, 1810: 164, entry 2-D). 176. Tmesipteris truncata R.Br. Bennett 121 (on same sheet as Bennett 122) ‘Lycopodioides truncata [. . .] Port Jackson [reverse] 2 Psilotum truncatum’ (Brown, 1810: 164, entry 2-J). FILICOPSIDA (TRUE FERNS) 177. Acrostichum aureum L. Bennett 2 Two labels: 1st reads ‘Acrostichum fraxinifolium prod 145 Arnhem South Bay Feb 6 1803’, Caledon Bay, Northern Territory (Burbidge, 1956); 2nd reads ‘Acrostichum fraxi- nifolium [. . .] Port II prod 145’, Port Clinton, Queensland (Burbidge, 1956). Brown was there in August 1802 (Vallance et a/., in prep.; Brown, 1810: 145, entry 2-T). 178. Adiantum aethiopicum L. Bennett 69 ‘Adiantum assimile Shoal Water Bay Keppel Bay prodr 155.’. A. aethiopicum of Beadle et al. (1972), perhaps Brown (1810: 155, entry 4). Brown, however, does not note a tropical specimen (“I”). 179. Adiantum aethiopicum L. Bennett 70 ‘Kangar|oo] Island’ (Brown, 1810: 155, entry 4-M). 180. Adzantum aethiopicum L. Bennett 71 ‘Adiantum ebeneum Port Jackson’ (Brown, 1810: 155, entry 4-J). 181. Adiantum formosum R.Br. Bennett 67 ‘Adiantum formosum prodr 155 Port Jackson’ (Brown, 1810: 155, entry 3-J). 182. Adiantum formosum R.Br. Bennett 68 ‘Adiantum cfr formosum Port Jackson [. . .]’ (Brown, 1810: 155, entry 3-J). 183. Adiantum hispidulum Sw. Bennett 64 ‘Adiantum hispidulum Shoal Water Bay prodr 155’ (Brown, 1810: 155, entry 2-T). 184. Adiantum hispidulum Sw. Bennett 65 ‘Adiantum [. . .] hispidulosa Port Jackson’ (Brown, 1810: 155, entry 2-J). PROC. LINN. SOC. N.S.W., 111 (2), 1989 86 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 185. Adiantum hispidulum Sw. Bennett 66 Three labels: 1st reads ‘Adiantum cfr A. hispidulum Port Jackson; 2nd reads ‘North rocks 8 August 1802’; (Brown, 1810: 155, entry 2-J). 3rd reads ‘Dove Dale Feb 1804 —’. 186. Arthropteris tenella (Forst.f.) J.Sm. Bennett 12 ‘Polypodium tenellium Forst prodr 147 Polypodium fimiculus rupibus & rare in loc umbrosis in rip[as] Grose’ (Brown, 1810: 147, entry 8-J). 187. Asplenium aethiopicum (Burm.f.) Bech. Bennett 33 (Syn. ‘Asplenium furcatum’ Thunb.) ‘Asplenium praemorsum prodr 150 rup umbrose: Grose’ (Brown, 1810: 150, entry 7-J). 188. Asplenium attenuatum R.Br. Unnumbered, but corresponds to Bennett register entry 25 No original label; secondary label reads ‘Nova Cambria Australis: Port Jackson R Brown (Brown, 1810: 150, entry 2-J). A duplicate of this in Kew Herbarium, as No 26, is from the ‘Grose’ River, Blue Mountains. 189. Asplenium bulbiferum Forst.f. Bennett 35 Two labels: Ist reads ‘Asplenium |. . .] laxum prodr 151’; 2nd reads ‘Asplenium laxum prodr 151 (Brown, 1810: 151, entry 10-D). 190. Asplenium flabellifolium Cay. Unnumbered, but corresponds to Bennett register 30 ‘Asplenium flabellifolium Van Diemens Land’ (Brown, 1810: 150, entry 6-D). 191. Asplenium flabellifolium Cay. Unnumbered but corresponds to Bennett register 31 ‘Asplenium flabellifolium Port Jackson’ (Brown, 1810: 150, entry 6-J). 192. Asplenium nidus L. Unnumbered but corresponds to Bennett register 25 (bis) ‘Asplenium nidus Port Jackson’ (Brown, 1810: 150, entry 1-J). 193. Asplenium nidus L. Unnumbered but corresponds to Bennett number 25 (bis) Three labels: Ist reads ‘Asplen. nidus North Coast’; 2nd reads ‘Asplenium nidus Linn prodr 150 Broad Sound West Peaked Hill In sylvis umbrosis ad radices arborum spec parasiticum” [cf. Brown, 1810: 150, entry 1-T]; 3rd reads ‘Grose 1804 (cf. Brown 1810: 150, entry 1-J). 194. Asplenium obtusatum Forst.f. Bennett 27 No original label; secondary label reads ‘Insula Van Diemen 1804 (probably Brown, 1810: 150, entry 3-D). 195. Asplenium paleaceum R.Br. Unnumbered but corresponds to Bennett register 29 No original label, secondary label reads ‘Nova Cambria australis in monte prope Broad Sound 1802 Robert Brown’ (cf. Brown, 1810: 150, entry 5-T). 196. Asplenium polyodon Forst.f. Unnumbered but probably corresponds to (Syn ‘A. falcatum’ Lam.) Bennett register 28 No original label; secondary label reads ‘Nova Cambria australis Port Jackson’ (perhaps corresponds with Brown, 1810: 150, entry 4-J). PROC. LINN. SOG. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 87 197. Azolla pinnata R.Br. Bennett 134 Two labels: 1st reads ‘Azolla pinnata Nob in stagnis prope Richmond in Noy Camb: durt’; 2nd reads ‘Azolla pinnata Nob in stagnes & fluv len: fluat Richmond Hawkesbury’ (Brown, 1810: 167, entry 1-J). Bauer painting 229 (BM (NH) Botany Library) is probably based on this plant or the specimen below. 198. Azolla pinnata R.Br. Bennett 135 ‘Azolla filiciloies [sic] pinnata Patersons River Oct 1804 (Brown, 1810: 167, entry 1-J). BM (NH) Bauer painting 229 perhaps based on this plant, but see above. 199. Azolla filiculoides Lam. var. rubra (R.Br.) Strasburger Bennett 136 ‘Azolla rubra Patersons River Oct 1804 (‘Azolla rubra’ of Brown, 1810: 167, entry 2-J). 200. Blechnum camfieldi Vindale Bennett 49 ‘Blechnum mutatum var. Port Jackson 8 Stegania procera prodr 153’ (‘Stegania procera of Brown, 1810: 153, entry 8-J). 201. Blechnum ambiguum (Presl.) Kaulf. ex C.Chr. Bennett 41 Two sheets, with identical labels ‘3 Blechnum laevigatum Prodr 152 Port Jackson’ (Brown, 1810: 152, entry 3-J). 202. Blechnum cartilagineum Sw. Bennett 39 ‘Blechnum paludosum Port Jackson [. . .] 1 Blechnum cartilagineum (Sw) prodr 152’ (cf. Brown, 1810: 152, entry 1-J). 203. Blechnum chambersi Tindale Bennett 42 ‘2 Stegania lanceolata prodr 152 Van Diemens Land’ (Brown, 1810: 152, entry 2-D). 204. Blechnum fluviatile (R.Br.) E. J. Lowe Bennett 43 ‘Pteris fluviatilis rivul alp Mont tabul. Derwent [. . .] 3 Stegania fluviatilis prodr 152’ (cf. Brown, 1810: 152, entry 3-D). 205. Blechnum indicum Burm.f. Bennett 40 ‘2 Blechnum striatum prodr 152 Port Jackson’ (Brown, 1810: 152, entry 2-J). 206. Blechnum minus (R.Br.) Ettingsh. Unnumbered but probably corresponds ‘Insula Van Diemen 1804 RB’ (Brown, 1810: 153, entry 7-D). to Bennett register 47 207. Blechnum nudum (Labill.) Mett. ex Luerss. Bennett 45 ‘Blechnum pteroides Port Dalrymple Jan 1804 5 Stegania nuda prodr 153’ (‘Stegania nuda’ of Brown, 1810: 153, 5-D). 208. Blechnum nudum (Labill.) Mett. ex Luerss. Bennett 46 ‘Blechnum ?discolour Osmunda discolor B&S Kings Id 6 Stegania falcata prodr 153 [. . .] (Stegania falcata’ of Brown, 1810: 153, entry 6-D). 209. Blechnum penna-marina (Poir.) Kuhn Bennett 44 ‘Pteris alpina desc cat Mont Tabul Derwent 4 Stegana alpina prodr 152 (‘Stegania alpina’ of Brown, 1810: 152, entry 4-D). PROC. LINN. SOC. N.S.W., 111 (2), 1989 88 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 210. Blechnum penna-marina ssp. alpina (R.Br.) Kuhn Unnumbered but probably correponds to Bennett register 44 ‘Insula Van Diemen in summitate Montis Tabularis 1804 RB Stegania alpina prodr [. . .| (probably another sheet of the above, cf. Brown, 1810: 152, entry 4-D). 211. Blechnum watts Tindale Bennett 48 ‘Blechnum mutatum desc Kings Island 8 Stegania procera prodr 153’ (Brown, 1810: 153, entry 8-D). 212. Blechnum watts Tindale Bennett 49 (bis) ‘Port Jackson 8 Stegania procera prodr 153’ (cf. “Stegania procera of Brown, 1810: 153, entry 8-J). The label bears an annotation by T. C. Chambers dated 6. x11. 1977 which reads ‘Maybe a hybrid but closest to Blechnum watts Tindale. 213. Botrychium australe R.Br. Bennett 119 ‘1 Botrychium australe prodr 164 Paramatta Port Jackson (Brown, 1810: 164, entry 1-J). 214. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 74 ‘Adiantoides crispa Kangar[oo] Id’. See Quirk et al. (1983: 510-3) regarding the relationship of Chezlanthes austrotenuifolia and C. tenuifolia. 215. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 76 ‘Adiantoides crispa Port Jackson (see above, and Quirk et al., 1983: 510-3). 216. Cheilanthes austrotenuifolia Quirk & Chambers Bennett 77 ‘Goose Id Bay South Coast Adiantoides crispa var’ (see above, and Quirk et al., 1983: D10=3)): 217. Cheilanthes brown (Desv.) Domin Bennett 5(a) ‘Arnhem South Bay Point U, FebY 6 1803 2 Notholaena villea prodr 146’ Mt. Caledon, Caledon Bay, Northern Territory (Burbidge, 1956; Vallance e¢ al., in prep.). ‘Notholaena vellea of Brown (1810: 146, entry 2‘T). Holotype (Quirk et al., 1983: SAB). 218. Chewlanthes browni (Desv.) Domin Bennett 5(b) ‘Shoalwater Bay Conical Pine Hill’ (Perhaps ‘Notholaena vellea of Brown, 1810: 146, entry 2-T). 219. Cheilanthes caudata R.Br. Bennett 78 ‘Pteris [struck through] Cheilanthes caudata prodr 156 Port II’ Port Clinton, Queens- land (Burbidge, 1956). Visited by Brown in August 1802 (Vallance ef al., in prep.). Brown (1810: 156, entry 2-T). Holotype (Quirk et al. 1983: 540-1). 220. Cheilanthes distans (R.Br.) Mett. Bennett 4(a) ‘Adiantoides palacea Port Jackson |. . .] Notholaena distans prodr 146’ (“Notholaena distans’ of Brown, 1810: 146, entry 1-J). Holotype (Quirk et a/., 1983: 526-9). 221. Chewlanthes lasiophylla Pichi-Serm. Bennett 4b ‘Adiantoides paleacea Inlet XII [. . .] Notholaena cfr N. distans’ At the Head of Spencer Gulf (Burbidge 1956), but Brown (1810: 146, entry 1) indicates PROG. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 89 Port Jackson as the source of the material. The Prodromus entry should have the ‘M’ annotation if Spencer Gulf is meant. See Quirk et al. (1983: 531-2) and Pichi-Sermolli (1951) regarding relationship with ‘Notholaena canescens. 222. Cheilanthes nudiuscula (R.Br.) T. Moore Bennett 60 ‘Pteris denudata v[ar] N Coast Id. y2’ Pibassoo’s Island, Northern Territory (Burbidge, 1956). ‘Pteris nudiuscula of Brown (1810: 155, entry 8-T). Holotype (Quirk e¢ al. , 1983: 537-40). 223. Cheilanthes tenuissima Bailey Bennett 61 ‘Adiantoides pubescens North Coast Island y2 &C’. Pibassoo’s Island, as above. 224. Cheilanthes tenuissima Bailey Bennett 3 ‘Acrostichum gracile [struck through] pteroides prodr 145 North Coast Islands 1803 RB’ (Brown 1810: 145, entry 3-IT). Type of ‘Acrostichum pteroides’ R.Br. (Quirk et al., 1983: Saco) 225. Cheilanthes tenuifolia (Burm.f.) Sw. Bennett 72 (Syn. ssp. queenslandica Domin., see remarks on relationship of C. austrotenuifolia and C. tenurfolia by Quirk et al. (1983: 506-13). ‘Adiantoides crispa var. North Coast Islands’ (Brown, 1810: 155, entry 1-T). 226. Cheilanthes sp.indet. Bennett 73 ‘Adiantoides crispa Port II’, Port Clinton area, Queensland (Burbidge, 1956). 227. Cheilanthes sp.indet. Bennett 75 ‘Adiantoides cicutaria Pteris [sic] Port IT’ 228. Christella dentata (Forsk.) Browns. & Jermy Bennett 21(a) ‘5 Nephrodium molle Prodr 149 [reverse reads] Plenosouris Polypod: molle Jacq. Grose’ (Brown, 1810: 149, entry 5-J). 229. Christella dentata (Forsk.) Browns. & Jermy Bennett 21(b) ‘Polypodium molle Jacq adrip: Patersons Hunter William Rivers det 1804 [reverse reads] 5 Nephrodium molle Prodr 149’ (Brown, 1810: 149, entry 5-J). 230. Ctenopterrs heterophylla (Labill.) Tindale Bennett 13 ‘Polypodium arbor prope in caulis Dicksonia Derwent 9 Polypodium grammitidis prodr 147 (Polypodium grammitidis’ of Brown, 1810: 147, entry 9-D). 231. Culcita dubia (R.Br.) Maxon Bennett 86 Two labels: 1st reads ‘Cfr D. cicutarian [struck through] Banks of Hunter’s River Nov 1804 4 Davallia dubia prodr 157’; 2nd reads ‘Davallia dubia Port Phillip’ (Brown, 1810: 157, entry 4-J and D). 232. Culcita dubia (R.Br.) Maxon Bennett 87 ‘4 Davallia dubia adripes fl. Patersons & Hunters Rivers Oct: Nov: 1804 (Brown, 1810: 157, entry 4-J). 233. Culcita dubia (R.Br.) Maxon Bennett 88 Two identical sheets “4 Davallia dubia Pt Jacksor (Brown, 1810: 157, entry 4-J). PROC. LINN. SOC. N.S.W., 111 (2), 1989 90 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 234. Culcita dubia (R.Br.) Maxon Bennett 89 ‘4 Davallia multiflora [struck through] dubia Port Jackson’ (Brown, 1810: 157, entry 4-J). 235. Culcita dubia (R.Br.) Maxon Bennett 90 ‘4 Davallia dubia prodr 157 Port Dalrymple Jan 1804 (Brown, 1810: 157, entry 4-D). 236. Cyathea australis (R.Br.) Domin Bennett 94 ‘Cyalloides Kings Island Basses Strait’ (Alsophila australis’ of Brown, 1810: entry 1-D). A tree fern, occurring in gullies near the sea. Lectotype (Tindale, 1956: 350). 237. Cyathea australis (R.Br.) Domin Bennett 95 ‘1 Alsophila australis prodr 158’ (probably ‘Alsophila australis’ of Brown, 1810: 158, entry 1-J). 238. Davallia pyxidata Cav. Bennett 84 ‘Davallia glycopoda P Jackson in rep (Brown, 1810: 156-7, entry 1-J). 239. Davallia pyxidata Cav. Bennett 85 Two sheets: Ist reads “Davallia pyxidata b Hunters River, Ash Island’; 2nd reads ‘Davallia glycopoda P Jackson in rup Grose [sic] (Brown, 1810: 156-7, entry 1-J). 240. Dennstaedtia davalliordes (R.Br.) T. Moore Bennett 93 ‘Davallia tenera Banks of Patersons River Oct: 1804 Dicksonia davallioides prodr 158 (Brown, 1810: 158, entry 2-J). 241. Diucksonia antarctica Labill. Bennett 92 Two sheets: Ist reads “Dicksonia australis Kings Island’; 2nd reads ‘Dicksonia antarctica Van Diemen [sic] Kings Island’ (second label relates to Brown, 1810: 157, entry 1-D. See also next). 242. Diucksonia antarctica Labill. Bennett 91 Two labels: Ist reads ‘Dicksonia australis Derwent; 2nd reads ‘Dicksonia antarctica prodr 157 Derwent’ (Brown, 1810: 157, entry 1-D). 243. Dicranopterts linearis (Burm.f.) L. Underw. Bennett 111 ‘Polypodium dichotomum Shoalwater bay passage |. . .] 8 Gleichenia Hermanni prodr 161 (Brown, 1810: 161, entry 8-1). 244. Diplazium australe (R.Br.) Wakef. Unnumbered, but probably corresponds to Bennett register entry 24 No original labels, but on back of sheet, not in Brown’s hand, ‘Nova cambria australe Port Jackson (perhaps ‘Allantodia tenera R.Br., cf. Brown, 1810: 149, entry 2-J). 245. Doodia aspera R.Br. Bennett 36 ‘Woodwardia aspera Banks of Hunters River frequent Oct-Novr 1804 Doodia aspera prodr 151 (Brown, 1810: 151, entry 1-J). 246. Doodia caudata (Cav.) R.Br. Bennett 37 ‘Woodwardia blechnoides sp5 [sic] Port Jackson [. ..] Doodia caudata prodr 151’ (Brown, 1810: 151, entry 3-J). PROG. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 91 247. Doodia caudata (Cav.) R.Br. Bennett 38 ‘Woodwardia Port Dalrymple Jan: 1804 (Brown, 1810: 151, entry 3-D). 248. Doryopterts concolor (Langsd. & Fisch.) Kuhn Bennett 62 ‘Pteris pedata prodr 155 Broad Sound in Montibus prope Upper Head’ (Brown, 1810: 155), cain O30). 249. Drynaria rigidula (Sw.) Bedd. Bennett 14a (but should be 15) ‘Polypodium anomalum Carpentaria Groote Eyland Frondes sterilis sumatae breviores latiores [reverse reads] Polypodium quercifolium B non Lin’ (Brown, 1810: 147, entry 10-1). 250. Drynaria sparsisora (Desv.)'T. Moore Bennett 14b (bis) 10 Polypodium quercifolium Prodr 147 Carpentaria Coen River (Brown, 1810: 147, entry 10-T). 251. Drynaria sparsisora (Desv.) T. Moore Bennett 14b (bis) ‘Polypodium Keppel Bay (Brown, 1810: 147, entry 10-T). 252. Gleichenia alpina R.Br. Bennett 105 ‘3 Gleichenia alpina desc prodr 160 Summitat Table Mountain Derwent’ (Brown, 1810: 161, entry 3-D). 253. Gleichenia dicarpa R.Br. Bennett 108 (Syns. ‘Gleichenia circinnata’ Sw., ‘Calymella circinnata (Sw.) Ching.?) ‘Gleichenia pubescens sp [. . .] P Dalrymple desc’ (possibly Brown, 1810: 161, entry 5-D). 254. Gleichenia microphylla R.Br. Bennett 106 (Syn. ‘Calymella circinnata’ (Sw.) Ching.?) ‘Gleichenia cfr speluncae Port Jackson’ (Brown, 1810: 161, entry 4-J). 255. Gleichenia microphylla R.Br. Bennett 107 Two labels: 1st reads “4 Gleichenia microphylla Prodr 161 Kings Island’; 2nd reads ‘Port Jackson’ (Brown, 1810: 161, entry 4-.D (label 1) and J (label 2) ). 256. Gleichenia microphylla R.Br. Unnumbered No original label: secondary label reads ‘New Holland Kents Group Dec 1803’ (Brown, 1810: 161, entry 4-D). 257. Gleichenia rupestris R.Br. Bennett 104 (Syn. ‘Calymella rupestris’ (R.Br.) Ching.) ‘Gleichenia austriaca desc Port Jackson [. . .] 2 Gleichenia rupestris Port Jackson prodr 160’ (Brown, 1810: 160, entry 2-J). 258. Gleichenia speluncae R.Br. Bennett 103 (Syn. ‘G. circinnata var.?, or ‘G. microphylla R.Br. °). ‘Gleichenia subsimplex desc rup [. . .| Port Jackson’ (Brown, 1810: 160, entry 1-J). PROG. LINN. SOC. N.S.W., 111 (2), 1989 92 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 259. Grammitis billardierr Willd. Bennett 6b &c Label 6b reads ‘Ceterach polypodiodes rupestris in later Mont Tabul Derwent Gram- mitis australis prodr 146’ (Brown, 1810: 146, entry 1-D); 6c reads ‘Grammitis australis prodr 146 Port Jackson 1804 RB’ (Brown, 1810: 146, entry 1-J). 260. Histiopterts incisa (Thunb.) J.Sm. Bennett 52 ‘3 Pteris vespertilionis P Jack[son and] Derwent (Brown, 1810: 154, entry 3-J). 261. Hustiopteris incisa (Thunb.) J.Sm. Bennett 54 ‘3 Pteris vespertilonis prodr 154 Kings Island’ (Brown, 1810: 154, entry 3-D). 262. Histiopterrs incisa (Thunb.) J.Sm. Bennett 55 ‘Pteris vespertilonis B Port Jackson’ (Brown, 1810: 154, entry 3-J). 263. Hymenophyllum cupressiforme Labill. Bennett 100 ‘Hymenophyllum tunbridgense prodr 159 Derwent’ (Brown, 1810: 159, entry 4-D). 264. Hymenophyllum flabellatum Labill. Bennett 98 ‘Hymenophyllum nitens prodr 159 Derwent (Brown, 1810: 159, entry 2-D). 265. Hypolepis rugosula (Labill.) J.Sm. Bennett 16a ‘12 Polypodium rugosulum Prod 147 [reverse reads] Polypodium affine Kings Island’ (Brown, 1810: 147, entry 12-D). 266. Lastreopsis decomposita (R.Br.) Tindale Bennett 22 ‘Pteridioides acuminata Polypodium Involuc renif. adrip. Hunters River Oct 1804 7 Nephrodium decompositum prodr 149’. (‘Nephrodium decompositum’ of Brown, 1810: 149, entry 6-J). Holotype of L. decomposita (cf. Tindale, 1965: 320-3). 267. Lastreopsis decomposita (R.Br.) Tindale Bennett 23 ‘Polypodium tenerum Broad Sound Peaked West Hill in sylvis umbrosis [. . .] 7 Nephro- dium decompositum [struck through] tenerum prodr 149’ (Nephrodium tenerum’ of Brown, 1810: 149, entry 7-T). 268. Lindsaea ensifolia Sw. Bennett 80 ‘Lindsaea lanceolata Prodr 156 [reverse reads] Lindsaea pleroides [‘Vittaria struck through] Port II Shoal Water Bay passage’ (Brown, 1810: 156, entry 2-T). This specimen has been determined by K. U. Kramer in 1965 as the ssp. agatz (Brack.) Kramer. 269. Lindsaea ensifolia Sw. Bennett 81 ‘2 Lindsaea lanceolata [@] [reverse reads] Lindsaea polymorpha North Coast Island Y,’ Arnhem Bay area, N.T. (Burbidge, 1956; Brown, 1810: 156, entry 2-T). Determined by K. U. Kramer in 1965 as the ssp. agai (Brack.) Kramer. 270. Lindsaea linearis Sw. Bennett 79 Two labels: 1st reads ‘Lindsaea recurifera Port Jackson [reverse reads] Lindsaea linearis Prodr 156’; 2nd reads ‘1 Lindsaea linearis’ (Brown, 1810: 156, entry 1-J, and Kramer and Tindale, 1976: 112-3). PROC. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 93 271. Lindsaea microphylla Sw. Bennett 83 Original label damaged, remaining part reads ‘[. . .]| Port Jackson’ (Brown, 1810: 156, entry 4-J, and Kramer and Tindale, 1976: 95-8). 272. Lindsaea media R.Br. Bennett 82 ‘Lindsaea irregularis North Coast Island y2 Feb 18-24 1803 Z Pibassoo’s Island (Burbidge, 1956). (Brown, 1810: 156, entry 3-I, and Kramer and Tindale, 1976: 98-101). 273. Lygodium flexuosum (L.) Sw. Unnumbered but probably corresponds to Bennett register entry 116 ‘Nova Hollandia or septen trionalis 1803 RB’ (Lygodium semibipinnatum’ of Brown, oO G2 entry 2-1 ): 274. Lygodium microphyllum (Cav.) R.Br. Unnumbered but probably corresponds (Syn. ‘Lygodium scandens (L.) Sw.) to Bennett register entry 115 No original label; secondary labels reads ‘Nova Cambria australis inter tropicum RB’ (Lygodium microphyllum’ of Brown, 1810: 162, entry 1-T). 275. Marsilea angustifolia R.Br. Bennett 141 ‘3. Marsilea angustifolium Carpentaria Island g Dec 25 1802’, Vanderlins Island (Burbidge, 1956; cf. Vallance et al., in prep.). (Brown, 1810: 167, entry 3-T). 276. Marsilea hirsuta R.Br. Bennett 138 ‘Marsilea hirsuta Carpentaria Island f, Bountiful Island (Burbidge, 1956). Brown was there on 3 December 1802 (Vallance et al. in prep.). (Brown, 1810: 167, entry 2-T). 277. Marsilea hirsuta R.Br. Bennett 139 ‘2 Marsilea [quadrifolia struck through] hirsuta Broad Sound in frales de propes scites temporebus inundatis (perhaps Brown, 1810: 167, entry 2-T). ‘Marsilia quadrifolia’ of Brown's Diary, collected on 15 September 1802 (Vallance et a/., in prep.). 278. Marsilea hirsuta R.Br. Bennett 140 ‘2 Marsilea villosa [struck through] hirsuta in |. . .] Port Jackson’ (Brown, 1810: 167, entry 2-]). 279. Marsilea mutica Mett. Bennett 137 (Syn. ‘M. brown’ R.Br., cf. Beadle e¢ al. (1972), in index.) ‘1 Marsilea quadrifolia Port Jackson’ (Brown, 1810: 167, entry 1-J). 280. Marsilea mutica Mett. Unnumbered Two labels: 1st reads ‘Nova Cambria australis Port Jackson R Br’; 2nd reads ‘Marsilia australis RB Marsilia quadrifolia prodr [blank] (probably Brown, 1810: 167, entry 1-J). 281. Mecodium australe (Willd.) Copel Bennett 99 ‘3 Hymenophyllum flabellatum [‘undulatum’ struck through] Labill prodr 159 Derwent’ (Hymenophyllum flabellatum’ of Brown, 1810: 159, entry 3-D). 282. Mecodium rarum (R.Br.) Copel. Bennett 97 Two labels: Ist reads ‘1 Hymenophyllum rarum prodr 159 Derwent; 2nd _ reads ‘Hymenophyllum rarum prodr |. . .] Van Diemans Island 1804 RB’ (Brown, 1810: 159, entry 1-D). PROG. LINN. SOG. N.S.W., 111 (2), 1989 94 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 283. Maicrosorum punctatum (L.) Copel. Bennett 10 ‘5 Polypodium irioides [‘punctatum’ struck through] Prodr 147 Keppel Bay Shoal Water Bay Broad Sound’. Another sheet of the same gathering has a secondary label which reads ‘Nova Cambria Australis intra tropicum 1802 RB’ (Brown, 1810: 147, entry 5-I). 284. Nephrolepis hirsutula (Forst.f.) C. Presl Bennett 20 ‘Nephrodium exaltatum prodr 148 Plendioides davilliastrum desc [“Davallioides’ struck through] Port II East Coast’. Port Clinton, Queensland (Burbidge, 1956), (Brown, 1810: 148, entry 2-T). 285. Ophioglossum costatum R.Br. | Bennett 118 ‘2 Ophioglossum costatum Arnhem N Bay Feb 14 1803 desc 55’ (Brown, 1810: 163, entry ZAM, 286. FPellaea falcata (R.Br.) Fee Bennett 59 ‘Kents Islands & Hawkesbury 6 Pteris [‘adiantoides’ struck through] falcata prodr 15¥ (‘Pteris falcata of Brown, 1810: 154, entry 6-J and D). 287. FPellaea paradoxa (R.Br.) Hook. Bennett 63 ‘Port Jackson 1 Adiantum paradoxum prodr 155’ (‘Adiantum paradoxum’ of Brown, 1810: 155, entry 1-J). 288. Phymatosorus diversifolia (Willd.) Pichi Serm. Bennett lla-c Three sheets: 1st (Bennett Ila) reads ‘6 Polypodium billarderi prodr 147 Port Dalrymple in umbrosis & truncus arbor Jan 1804’; 2nd reads (Bennett 11b) “‘Polypodium poly- morphum var Port Jackson’; 3rd (Bennett 11c) reads ‘Kents Group Bass Strait 7 Poly- podium [‘lycopodioides’ struck through] Billardieri prodr 147’ ((Polypodium billardiery of Brown, 1810: 147, entry 7-D). The indication would appear to be that the Port Jackson specimen did not feature in the description. 289. Platycerrum bifurcatum (Cavy.) C.Chr. Bennett 1 Two labels: Ist reads ‘Acrostichum alcicorne prodr pl45 Port Jackson 1802-5 Rocks & ‘Trees’; 2nd reads ‘1 Acrostichum alcicorne prodr 145’ (Brown, 1810: 145, entry 1-J). 290. Platyzoma microphyllum R.Br. Bennett 101 (?Syn. ‘Gleichenia platyzoma’ F. Muell.) ‘Platyzoma microphylla Carpentaria Island h’ (North Island cf. Burbidge 1956) (Brown, 1810: 160, entry 1-T). 291. Platyzoma microphyllum R.Br. Bennett 102 (? Syn. ‘G. platyzoma F. Muell.) ‘Acrostichum? Gleichenia? Facing Island Port I’ (Queensland, cf. Brown, 1810: 160, entry 1-I). 292. Pleurosorus rutiflorus (R.Br.) Fee Bennett 7 ‘Derwent Risdon 2 Grammitis rutaefolia prodr 146’. Another specimen on this sheet bears the secondary locality ‘Nova Hollandia ora australis Kangaroo Island 1802 RB’ (probably ‘Grammitis rutaefolia’ of Brown, 1810: 146, entry 2-D). PROC. LINN. SOC. N.S.W,, 111 (2), 1989 E. W. GROVES AND D. T. MOORE 5 293. Polyphlebrum venosum (R.Br.) Copel. Bennett 96 ‘Derwent 1 Trichomanes [‘repandum’ struck through] venosum prodr 159’ (Brown, 1810: 159, entry 1-D). 294. Polystichum australiense Vindale Bennett 18 No original label; secondary annotation reads ‘Derwent’. Separated out from material of Bennett 17 (see below) and assigned by Bennett’s workers to No 18. 295. FPolystichum proliferum (R.Br.) C. Presl Bennett 17 ‘Aspidium proliferum prodr 147 desc Port Jackson [and] Derwent’ (Brown 1810: 147-8, entry 1-D). Lectotype (Tindale, 1961: 50). 296. Pteridium esculentum (Forst.f.) Cockayne Bennett 57 ‘Pteris esculenta var P Jackson’ (Brown, 1810: 154, entry 5-J). 297. Pteridium esculentum (Forst.f.) Cockayne Bennett 58 ‘D Pteris esculenta prodr 154 Derwent (Brown, 1810: 154, entry 5-D). 298. Pteris tremula R.Br. Bennett 53 ‘Grose’ (Brown, 1810: 154, entry 4-J). 299. Pteris tremula R.Br. Unnumbered but corresponds to Bennett register entry 56 ‘Pteris tremula prodr 154 Port Jackson’ (Brown, 1810: 154, entry 4-J). 300. Pteris umbrosa R.Br. Bennett 51 ‘Pteris umbrosa First branch of the Grose Pteris’ (Brown, 1810: 154, entry 1-J). 301. Pyrrosia confluens (R.Br.) Ching Bennett 9 ‘Pteroides varia [struck through] Polypodium confluens prodr 146 cfr Plenopilo- selloides ad rip Hunters & Patersons Rivers Oct: Nov 180¥ (‘Polypodium confluens’ of Brown, 1810: 146, entry 3-J). 302. Pyrrosia rupestris (R.Br.) Ching Bennett 8 ‘Polypodium rupestre prodr 146 [reverse] Polypodium serpens B & S Forst Prod: Vahl Symb Port Jackson’ (‘Polypodium rupestris’ of Brown, 1810: 146, entry 2-J). 303. Rumohra adiantiformis (Forst.f.) Ching Bennett 19 ‘2 Aspidium coriaceum prodr 148 Polypodium adiantiforme Forst prod Derwent adras montis (‘Aspidium coriaceum of Brown, 1810: 148, entry 2-D). 304. Schizaea bifida Willd. Bennett 113 Two labels: 1st reads; ‘3 Schizaea furcata [struck through] bifida prodr 162 Port II’; 2nd reads ‘Schizaea furcata Richmond’ (Brown, 1810: 162, entry 3-J. The provenance annotation is J and D; perhaps T and J are meant). 305. Schizaea bifida Willd. Unnumbered — Dryander duplicate, see introduction. ‘Acrostichum pectinastordes Nova Cambria australis Port Jackson’. 306. Schizaea dichotoma (L.) J.Sm. Bennett 114 ‘Schizaea dichotoma 4 Schizaea aspera scabr cfr S dichotom: prodr 162’ (Brown, 1810: 162, entry 4-T). PROG. LINN. SOG. N.S.W,, 111 (2), 1989 96 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 307. Schizaea fistulosa Labill. Unnumbered “Terra Diemam (‘Schizaea fistulosa of Brown, 1810: 162, entry 2-D). 308. Schizaea fistulosa Labill. Unnumbered (Syn. ‘Microschizaea fistulosa’ (Labill.) Reed). ‘Schizaea fistula[. . .]. Locality probably Tasmania as above (cf. Brown, 1810: 162, entry eID), 309. Schizaea rupestris R.Br. Bennett 112 ‘Schizaea rupestris prodr 162 desc 25 nep |. . .| Port Jackson’ (Brown, 1810: 162, entry 1-J). 310. Stucherus flabellatus (R.Br.) St. John Bennett 109 ‘Port Jackson 6 Gleichenia [‘prolifer’ struck through] flabellata prodr 161 (‘Gleichenia flabellata of Brown, 1810: 161, entry 6-J). 311. Sticherus tener (R.Br.) Ching Bennett 110 ‘7 Gleichenia tenera prodr 161 in rup. hum ad rip rivul alpin Mon|tis] Tabul{aris]’ (‘Gleichenia tenera of Brown, 1810: 161, entry 7-D). 312. ‘odea barbara (L.) T. Moore Bennett 117 ‘Osmunda barbara prodr 163 Port Jackson’ (Brown, 1810: 163, entry 1-J). 313. Vittaria elongata Sw. Bennett 50 ‘Vittaria acostata Broad Sound Peaked West Hill parasit: super Asplen|ium] nidus’ (Brown, 1810: 153, entry 1-T). CYCADOPHYTES Cycadaceae 314. Cycas angulata R.Br. Bennett 3106 (pro parte) No original label. According to an annotation by J. R. Maconochie of 18 November 1976, the specimen is probably from Bountiful Island, Gulf of Carpentaria, and the “I” entry confirms the tropical provenance (Brown, 1810: 348, entry 2-T). In his Diary (Vallance et al., in prep.) Brown does not mention a cycad being collected from Bountiful Island but Good mentions a ‘Cycas circinalis’ which his editor (Edwards, 1981: 104) identified as C. media R.Br. But we believe that C. angulata is correct. Bauer painting 159/60, Botany Library, BM (NH), is probably based on this plant. 315. Cycas media R.Br. Bennett 3106 (pro parte) No original label. According to an annotation by J. R. Maconochie of 18 November 1976, the specimen is probably from Cumberland Islands, Queensland. The “IT” entry (Brown, 1810: 348, entry 1-I) confirms this (cf. Mabberley, 1985: plate 4). Certainly Brown collected a specimen of ‘Cycas circinalis’ at Calder Island (Cumberland Islands) on 16 October 1802 which could be this specimen. BM (NH) Bauer painting 157/8 is probably based on this plant (Vallance e¢ al., in prep.). CONIFEROPHYTES Araucariaceae 316. Araucaria cunninghami Ait. Bennett 3115 PROG. LINN. SOG. N.S.W.,, 111 (2), 1989 E. W. GROVES AND D. T. MOORE 97 ‘Araucaria cunninghamii Port Bowen 1802 RB’ (probably the specimen collected 21 August 1802 according to Brown’s Diary (Vallance et al., in prep). Cupressaceae 317. Callitris gracilis R. Baker Bennett 3108 (Syn. ‘Frenela robusta’ A. Cunn.) Two labels: 1st reads ‘Callitris glauca Nob Mount Brown Spencers Gulf 1802 RB’; 2nd reads ‘Genus Conifer Inlet XII South C’. Gathered from Mount Brown, 10-11 March 1802 (cf. Vallance et a/., in prep.). 318. Callitris gracilis R. Baker Bennett 3109 (Syns ‘Frenela robusta’ A. Cunn., ‘Callitris propinqua’ R.Br.) ‘Callitris propinqua Nob Kanguroo Island 1802 RB’ Gathered March 1802 (cf. Vallance et al., in prep.). 319. Callitris muellert (Parl.) F. Muell. Bennett 3112 (Syn. ‘Frenela mueller’ Parl., cf. Garden, 1956: 383). ‘Callitris Nova Holland loc incerta. 320. Callitris rhomboidea R.Br. ex A. & L. C. Rich Bennett 3107 (Syn. Holotype of ‘Callitris ventenati’ Mirb., Garden, 1956: 386) ‘Callitris ventenati1 Nob Port Jackson 1804-5” 321. Callitris rhomboidea R.Br. ex A. & L. C. Rich Bennett 3114 (Syn. ‘Callitris australis’, cf. Garden, 1956: 386). ‘Genus Coniferar: adripas saxosus fluvii Cataract River Port Dalrymple Jan 10 1804 [. . .] Callitris australis’ 322. Callitris tuberculata R.Br. ex R. T. Baker & H. G. Smith Bennett 3110 (Syn. ‘Frenela robusta var. verrucosa, also ‘Callitris preiss1’ Miq.). Two labels: Ist reads ‘Middle Island of Goose Island Bay 1802 RB’; 2nd reads ‘Genus Conif Bay II South Coast 1802 Jan 12’. Gathered January 1802 from the Archipelago of the Recherche (cf. Willis, 1959: 99; and Vallance et al., in prep.). 323. Callitris sp.indet. Bennett 3111 ‘Port Jackson. 324. Callitris columellaris F. Muell. Bennett 3113 ‘Nov gen Conifer Sandy Cape Arbor Mediocris cortice ramoso crasso’. Collected 30 July 1802, Vallance et al. (in prep.). Podocarpaceae 325. Lagarostrobos franklini (Hook.t.) C. J. Quinn Bennett 3120 ‘Huon Pine’. No locality given on original label. This taxon sensu Quinn (1982). 326. Phyllocladus aspleniifolius (Labill.) Hook.f. Bennett 3119 (Syn. ‘Podocarpus asplenifolia’ Labill.). ‘Derwent [. . .] Podocarpus asplenifolia. PROG. LINN. SOG. N.S.W., 111 (2), 1989 98 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS 327. Podocarpus elatus R.Br. ex Endl. Bennett 3117 Two labels: Ist reads “Taxus cfr T elongatum Hunters River; 2nd reads “Iaxus cfr T elongatum In sylvis prope ripas fluvio Paterson A A Ash Island Insulas que adjacent Cabbage tree swamp & C Desc In mscr N C No95’. Probably gathered 14 October 1804 (cf. Vallance et al. in prep.). 328. Podocarpus lawrence: Hook.f. Bennett 3116 (‘Podocarpus alpinus’ of de Laubenfels, 1985). “Taxus emersa Nol20 desc Misc fl Derwent In Summitate saxosa Montis Tabularis prope fluio: Derwent Apr: Mai 180. 329. Podocarpus spinulosus (Sm.) R.Br. ex Mirb. Bennett 3118 No original label. INDEX Poorly-localized material is unindexed, but material from the south coast is given below. This is broadly equivalent to Brown’s (1810) annotation ‘M’: 3, PO, 4), 13, 20, 22, Bae, DD, BO=T, 42-3), 42], DD 13), GH GO, IZZ, WA) 72, UTS), Zil4e, SUS and 322. Material from Tasmania, and Bass Strait, approximately equivalent to Brown’s (1810) ‘D’ annotation: Dy 4, Dy Il, WIG), Zl, ZH, HlleB2,. HAs, DS, Gel, 43, HS=Hil, DS), DY, OA, GIS, /le3, 13-9, O7/, OY. GHG, 100, WOSabL, NSD, Wl7/3, 20225), IOS, W378, Wal, Wa 3=59, ID /=3,, IOW04, loo=7, WOME, W7/H,, USV-GO. Ws, AOI-O42, AVO=bl, ZHO=l, AIH=O, Z4wle4o2,, Wei), VU=3, YO, ZO, VOS= 5, ZMleZ, ZO, 2HS, 22-3, 299, 27, 303, 307-8, Dbl, BAN, BAG eral HAS: Material from New South Wales, equivalent to Brown’s (1810) ‘J’ annotation: QI-D, WOIO4, WG-29, UBS, WA), We, WHO, HOS, Wers,, W7Sie7/4s, WO, MOA, WSHecks, I=}, IQG-20Z, AVS, AIDA, VAY, ZAS=2Y), VBL=S4e, ZDI-B), VAL), WEA, VIO), BOL, AEG, Z/0= Mi, 2TS-80, 237, ZE9, 296, ZVS-3OZ, SO, SVQ, Si2, 320, S23 aval S27. Tropical material, equivalent to Brown’s (1810) “TI” annotation: Di), 29), 48, BO), 82, 84, W/T=78, MSS, DD, ZiT, LAA, ZHAaVS), 243}, ZAG-Gil, VOT, AOS=59), DIDI Ny POSBO, 2YO-DL, BOL, BOO, BBall, BLA? aanel B27. INDEX OF BENNETT NUMBERS vs GROVES & MOORE NUMBERS Wineee aie In lskernneitt mvtonl serge (3, Sa, Bis, 440), HO), Ges, GY, IG, WS), iil, ISS (| =25)]). ISO SHO), WN Sail, i225], Io [S245], 200 ||P Seer], Zl) ease ares |p 24], 256, 273 (2? =16), 274 [2=155], 280, 299 |=56], 305, 307 and 308. Possibly equivalent Bennett numbers are given in brackets — see text for each such entry. For Groves & Moore numbers marked with an asterisk (*) there is no Bennett number. Bennett Groves & Moore Bennett Groves& Moore Bennett Groves & Moore 1 289 9 301 19 508 2 a 10 283 20 284 3) 224 lla-c 288 2la 228 4a 220 12 186 21b 229 4b Pa 13 230 BD 266 al 27, allo) 249 BS 267 5b 218 14b BIO) Ce LHL 24 244* 6b &c 259 l6a 265 25 188, 192 & 193* 7 292 17 295 Di 194 8 302 18 294 28 196* PROG. LINN. SOC. N.S.W., 111 (2), 1989 E. W. GROVES AND D. T. MOORE 99 Bennett Groves & Moore Bennett Groves & Moore Bennett Groves & Moore 29 Is 81 269 131 168 30 tO Or 82 272 132 173 31 Oe 83 ZH 133 172 33 187 84 238 13)4¢ 197 35 189 85 L39) 139 198 36 245 86 POI 136 199 on 246 87 23y2 137 DUS 38 247 88 D9) 138 276 39 202 89 Dove 139 ZU 40 205 90 235 140 278 41 201 Dil 242 141 ZAS 42 203 92 Deel 157 39 43 204 93 240 158 41 44 209 & 210* 94 236 159 aes 45 207 95 ZU 161 44 46 208 96 2S)3} 162 45 ALT 206* a 282 163 46 48 211 & 212 98 264 166 3 49 200 yg 281 167 5 50 ols 100 263 168 6 51 300 101 290 169 7 oy 260 102 291 170 10 53 298 103 258 171 9) 54 261 104 257 UZ 4 55 262 105 DDD 173 2 56 DS ie 106 DO: 174 11 5/) 296 107 Dd)5) 7 32 58 ZS)i/ 108 ZS 176 69 a9) 286 109 310 177 a) 60 222 110 311 179 18 61 LS 111 243 181 2, ds} we wee 62 248 112 309 18la 15 63 287 113 304 182 a2 64 183 114 306 193 60 65 184 115? Dae 195 53 66 185 116? Duss 203 16 67 181 117 312 205 1 68 182 118 285 206 68 69 178 119 213 208 20 70 7) 120 174 209 22 ML 180 121 176 210 42 Ue 229 122 175 211 2 73 226 123 169 PS 7 74 Dist 124 170 214 21 75 2h 125 171 215 48 76 215 126 163 216 47 el 216 127 164 217 oy 78 ZAG 128 165 218 27 19 270 129 166 219 Uae 80 268 130 167 221 25 PROG. LINN. SOC. N.S.W., 111 (2), 1989 100 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS Bennett Groves & Moore Bennett Groves & Moore Bennett Groves & Moore 923 28 984 150 539 100 994 26 985 157 540 116 & 119 995 49 986 160 542 149 226 19 287 155 543 111 BAT 3D) 288 oul 544 128 228 50 289 152 545 134 229 31 290 144 546 97 ie Sr t Soecie 933 78 993 143 i ae 235 74 294 ie a iz 936 63 995 14 See! 240 54 296 149 eee eee 244 16 501 129 I! tee 245 55 502 124 399 133 249 li 504 99 556 112 250 57 507) 104 558 137 951 30 & 71 511 93 560 102 952 72 515 107 961 101 253 64 516 118 963 96 254 61 Ry 130 565 109 957 65 518 139 & 140 566 110 959 62 519 126 567 120 260 67 520 136 569 103 261 80 521 114 3106 314 & 315 963 66 522 98 3107 320 264 56 HLS 138 3108 7 266 75 S28 131 3109 318 267 79 525 141 3110 399 268 73 526 105 & 132 3111 393 969 59 527 91 3119 319 971 51 528 135 ove AO. 972 58 530 90 tnd < 276 83 531 89 i 277 81 532 85 ce Ve 279 82 533 86 3116 328 280 162 534 87 & 88 S117 327 281 155 536 115 3118 329 282 154 537 113 3119 326 983 158 538 117 3120 395 ACKNOWLEDGEMENTS A work of this nature could not have been undertaken without the assistance of specialist taxonomists and curators. Consequently in the British Museum (Natural History) we wish to record our thanks to Miss J. M. Camus, Dr D. J. Galloway, Mr L. Ellis, Dr A. J. Harrington, Dr C. R. Hill, Mr S. I. Honey, Mrs L. M. Irvine, Mr J. H. Price, and Mr I. Tittley. We also thank the Botany Librarian, Mrs C. V. Ellwood, for access to the Brown slip catalogue and Mr J. F. M. Cannon, Keeper of Botany, also for access to the collections in his charge. We owe a considerable debt to our Australian friends, particularly to Mr K. Hill PROG. LINN. SOG. N.S.W., 111 (2), 1989 E. W. GROVES AND D. ‘T. MOORE 101 and Mrs Karen Wilson of the Royal Botanic Gardens, Sydney, and also Professor T. G. Vallance of the University of Sydney. We thank Dr I. R. Price, James Cook University, Queensland, for information on Brown’s algae, and Dr M. A. Clements of the Australian National Botanic Gardens, Canberra, for helpful discussion. References AGARDH, C. A., 1823-28. — Species Algarum rite cognitae cum synonymis. Gryphiswaldiae: sumtibus E. Mavritil. 2 vols. BEADLE, N. C. W., Evans, O. D., CAROLIN, R. C., and ViINbDALE, M. D., 1972. — Flora of the Sydney Region. Sydney: A. H. & A. W. Reed. BENTHAM, G., and MUELLER, F. VON, 1863-1878. — Flora Australiensis: A description of the plants of the Australian Territory. London: Lovell Reeve & Co. 7 vols. Bisby, P., 1951. — Parmelia australiensis Cromb. Vict. Nat. 67: 186. BRITTEN, J., 1904. — R. Brown’s list of Madeira plants. J. Bot., Lond. 42: 1-8, 39-46, 175-182 and 197-200. BROWN, R., 1810. — Prodromus Florae Novae Hollandiae et Insulae Van-Diemen, exhibens characteres plantarum quas annis 1802-1805 per oras utriusque insulae collegit et descripsit Robertus Brown; . . . London: ‘Typis Richardi Taylor ct Soci. The work, with the Supplementum Primum 1830, was reprinted in facsimile 1960 — Weinheim, Codicote and New York: Engelmann, Wheldon & Wesley Ltd and Hafnter. —— , 1811. — Some observations on the parts of fructification in mosses; with characters and descriptions of two new gencra of that order. Zrans Linn. Soc. Lond. 10: 312-324. ——, 1826. — On the structure of the female flower of Cycadcac and Coniferac. /n KING, P. P., Narrative of a survey of the intertropical and western coasts of Australia performed between the years 1818 and 1822. Vol. 2: 554- 565. London: John Murray. BuURBIDGE, N. T., 1956. — Robert Brown’s collecting localities. Proc. Linn. Soc. N.S.W. 80: 229-233. CLEMENTS, M. A., 1983. — Notes on the content of John Lindley’s Orchid herbarium. Orchadian 7(10): 241-245. CromMBIE, J. M., 1879. — Enumeration of Australian lichens in Herb. Robert Brown (Brit. Mus.) with descriptions of new species. J. Linn. Soc. Lond. (Bot.) 17: 390-401. Drcetius, G., 1974. — The lichen genus Collema with special reference to the extra-European species. Acta Univ. Upsaliensis, Symbol. Bot. XX. 2. Dr LAUBENFELS, D. J., 1985. — A taxonomic revision of the genus Podocarpus. Blumea 30: 251-278. Epwarps, P. I., 1976. — ‘Robert Brown (1773-1858) and the natural history of Matthew Flinders voyage in H.M.S. Investigator 1801- 1805. J. Soc. Biblphy. nat. Hist. 7: 385-407. , ed., 1981. — The Journal of Peter Good: Gardener on Matthew Flinders’ voyage to Terra Australis 1801-03. Bull. Br. Mus. nat. Hist. (hast. Ser.) 9: 1-213. ELIX, J. A., JOHNSON, J., and ARMSTRONG, P. M., 1986. — A revision of the lichen genus Xanthoparmelia in Australia. Bull. Br. Mus. nat. Hist. (bot. Ser.) 15: 163-362. FILSON, R. B., 1981. — A revision of the lichen genus Cladia Ny]. J. Hattori bot. Lab. 49: 1-75. FLINDERS, M., 1814. — A voyage to Terra Australis... in the years 1801, 1802 and 1803, in His Majesty’ ship the Investigator. . . London: G. & W. Nicol. 2-vols& Atlas. FORBES, H. O., 1885. — A naturalist’s wanderings in the Eastern Archipelago, from 1878 to 1863. Appendix V1: 497- 523. London: Sampson Low. GaLLoway, D. J., 1985. — Flora of New Zealand Lichens. Wellington: Government Printer. GARDEN, J., 1956. — A revision of the genus Callitris Vent. Contr. N.S.W. Herb. 2: 363-392. Gray, J. E., 1866. — On Anadyomene and Microrictyon, with the description of three new gencra, discovered by Menzies in the Gulf of Mexico. /. Bot., Lond. 4: 41-51, 65-72 and 291-293. GorrscHE, C. M., LINDENBERG, J. B. G., and NEES VON ESENBECK, C. G., 1844-7. — Synopsis Hepaticarum. Hamburegi: sumtibus Mcissnerianis. Groves, E. W. and Moore, D.'T., 1986. — Preparation of a first listing of the plants collected in Australia by Robert Brown (1773-1859) during the Flinders voyage of 1801-1805. News. Aust. Spstem. Bot. Soc. (December 1986): 4-5. JOHNSON, L. A. S., 1959. — The families of cycads and the Zamiaccac of Australia. Proc. Linn. Soc. N.S.W. 84: 64-117. KANTVILAS, G., 1983. — A brict history of lichenology in Tasmania. Pap. Proc. Roy. Soc. Tasm. 117: 41-31. KRAMER, K. U., and TINDALE, M. D., 1976. = The Lindsacoid ferns of the Old World. VII. Australia and New Zealand. Yélopea 1: 91-128. LINDLEY, J., 1844. — Review of J. D. HOOKER on “The botany of the Antarctic voyage of H.M?s discovery ships Erebus and ‘Icrror, in the years 1839-1843". Gdnrs Chron. 27: 446. MABBERLEY, D. J., 1985. — Jupiter Botanicus: Robert Brown of the British Museum. Braunschweig and London: J. Cramer and British Museum (Natural History). PROG. LINN. SOG. N.S.W., 111 (2), 1989 102 ROBERT BROWN’S AUSTRALIAN CRYPTOGAMS AND GYMNOSPERMS Murray, G., 1904. — The Department of Botany. In The History of the Collections contained in the Natural History Departments of the British Museum. Vol. 1. London: British Museum. Norst, M. J., (in press). — Ferdinand Bauer. The Australian natural history drawings. London: British Museum (Natural History). PICHI-SERMOLLL, R. E. G., 1951. — Notes on some Australian ferns. Webbia 8: 205-211. POWELL, M., and MORLEY, B., 1976. — The Robert Brown material in the National Herbarium, Glasnevin, Dublin. Contr. Nat. Bot. Gdns. Ireland, Glasnevin, Number 1. Prick, J. H., 1984. — Bibliographic notes on works concerning the Algae. V: A note on aspects of the Fucz. . . (Dawson Turner, 1807-19). Archiv. nat. Hist. 11: 431-442. QUINN, C. J., 1982. — Taxonomy of Dacrydium Sol. ex Lamb. emend. de Laub. (Podocarpaceac). Aust. J. Bot. 30: 311-320. Quirk, H., CHAMBERS, T. C., and REGAN, M., 1983. — The fern genus Cheilanthes in Australia. Aust. J. Bot. 31: 501-553. ROURKE, J. P., 1974. — Robert Brown at the Cape of Good Hope. /. S. Afr. Bot. 40: 47-60. Scott, G. A. M., 1985. — Southern Australian Liverworts. Austr, Flora G Fauna Ser. no. 2. Canberra: Australian Government Publishing Service. , and STONE, I. G., 1976. — The mosses of southern Australia. San Francisco: Academic Press. STEARN, W. T., 1960a. — An introduction to Robert Brown’s Prodromus Flora Novae Hollandiae. Weinheim and Codicote: Engelmann and Wheldon & Wesley Ltd. — ., 1960b. — Franz & Ferdinand Bauer, masters of botanical illustration. Endeavour 19: 27-35. ——, 1981. — The Natural History Museum at South Kensington. London: Heinemann and British Muscum (Natural History). , and BLunT, W., 1976. — The Australian Flower Paintings of Ferdinand Bauer. London: The Basilisk Press. TINDALE, M. D., 1956. — Cyatheaceae of Australia. Contr. N.S.W. Herb. 2: 327-61. —., 1961. — Pteridophyta of south-eastern Australia. Number 211, Aspidiaceae. Contr. N.S.W. Herb. Flora Ser, 47-78. ——, 1965. — A monograph of the genus Lastreopteris Ching. Contr. N.S.W. Herb. 3: 249-3. TURNER, D., 1807-1819. — Fucz; or coloured figures and descriptions of the plants referred by botanists to the genus Fucus. London: John & Arthur Arch. 4 vols. VALLANCE, T. G., and Moore, D. T., 1982. — Geological aspects of the voyage of H.M.S. Investigator in Australian waters, 1801-5. Bull. Br. Mus. nat. Hist. (hist. Ser.) 10: 1-43. ——., GROVES, E. W., and Moore, D. T., eds, (in preparation). — Nature’ Investigator: The diary of Robert Brown in Australia 1801-5. WALKER, F. J., 1985. — The lichen genus Usnea subgenus Neuropogon. Bull. Br. Mus. nat. Hist. (bot. Ser.) 13: 1-130. WILLIS, J. H., 1959. — Plants of the Recherche Archipelago, W.A. Muelleria 1: 97-101. WoMERSLEY, H. B. S., 1967. — A critical survey of the marine algae of southern Australia. Aust. J. Bot. 15: 189-270. ——.,, 1984. — The marine benthic Flora of South Australia. Part 1. Adelaide: Government Printer. —— , 1987. — The marine benthic Flora of South Australia. Part 11. Adelaide: Government Printer. WOobD, R. D., 1965. — Monograph of the Characeae. Weinheim: J. Cramer. WoopbwarbD, B. B., 1904. — Libraries. In The History of the Collections contained in the Natural History Departments of the British Museum. Vol. 1. London: British Museum. PROC. LINN. SOC. N.S.W., 111 (2), 1989 The Lichens of Norfolk Island 1: Introduction and the Family Parmeliaceae JOHN A. ELIX and HEINAR STREIMANN ELIx, J. A., & STREIMANN, H. The lichens of Norfolk Island. 1: Introduction and the family Parmeliaceac. Proc. Linn. Soc. N.S.W. 111 (2), 1989: 103-121. The historical records of lichens of the Norfolk Island complex are discussed and synonymy given. The 16 species (from 7 genera) of the Parmeliaceae which currently grow there are examined critically. Each species is described fully (including chemistry) and its distribution is recorded. The endemic Flavoparmelia norfolkensis Elix & Streim. is described as new to science. All 16 species on these geologically recent, oceanic islands possessed vegetative dispores, attesting to the efficiency of this method of reproduction. John A. Elix, Department of Chemistry, The Faculties, Australian National University, G.P-O. Box 4, Canberra, Australia 2601, and Heinar Streimann, Cryptogamic Herbarium, Australian National Botanic Gardens, G.P.O. Box 1777, Canberra, Australia 2601; manuscript receved 23 November 1988, accepted for publication 15 February 1989. KEY WORDS Flavoparmelia norfolkensis Elix & Streim., Neofuscelia verrucella (Essl.) Essl., Paraparmelia scotophylla (Kurok.) Elix & Johnston, Parmelia erumpens Kurok., Parmelinopsis spumosa (Asahina) Elix & Hale, Farmotrema austrocetratum Elix & Johnston, Parmotrema chinense (Osbeck) Hale & Ahti, Parmotrema crinitum (Ach.) Choisy, Parmotrema cristiferum (Taylor) Hale, Parmotrema gardnen (Dodge) Sérusiaux, Parmotrema rampoddense (Nyl.) Hale, Parmotrema reticulatum (Taylor) Choisy, Parmotrema sancti-angelu (Lynge) Hale, Parmotrema tinctorum (Despr. ex Nyl.) Hale, Xanthoparmelia amplexula (Stirton) Ehx & Johnston, Xanthoparmelia australasica D. Gall., Parmeliaceae, chemotaxonomy, Norfolk Island, lichens. INTRODUCTION The Norfolk Island complex (including Norfolk, Phillip and Nepean Islands with the satellite islands) is an isolated volcanic outcrop in the South Pacific Ocean between longitudes 167°55’ and 168°00’E and latitudes 28°59’ and 29°08’S. The islands lie on a narrow, steep-sided submarine ridge (the Norfolk Ridge) which extends from New Caledonia to New Zealand (Jones and McDougall, 1973). The closest land is New Caledonia, 670 kilometres to the north. Norfolk Island les approximately 1610 kilometres north-east of Sydney and_1100 kilometres north-north-west of Auckland (Eig): The islands are an Australian territory: Norfolk Island (the largest) is 8 kilometres long and 6 kilometres wide; Phillip Island is 2 kilometres long and 1 kilometre wide, while Nepean Island is approximately 500 metres long and 300 metres wide (Figure 2). Norfolk and Phillip Islands have similar geology: both are almost completely volcanic in origin. Olivine basalt lavas predominate on Norfolk Island, with basaltic tuffs relatively more common on Phillip Island. This volcanic activity occurred 2.3-3.2 million years ago, during the Pliocene epoch (Jones and McDougall, 1973). Nepean Island and part of Norfolk Island (near Kingston) consist of coarse marine calcareous rock of late Pleistocene origin (Ovington, 1984). The highest point on Norfolk Island is Mt Bates, which is 319 metres above sea level. Mt Pitt is slightly lower (318m), but these peaks are only 600 metres apart and together form the summit complex. The southern half and north-west corner of the island are gullied plateaux 90-120 metres a.s.]. and are distinct from the Mt Pitt — Mt Bates summit area. Most of the coastline, except for a length on the southern coast (Kingston area) and several bays, consists of cliffs up to 140m high (Jones and McDougall, 1973; Ovington, 1984). PROC. LINN. SOC. N.S.W., 111 (2), 1989 104 LICHENS OF NORFOLK ISLAND 1 Fig. 1. Location of Norfolk Island. Phillip Island is even more rugged and precipitous, rising to its highest point at Jacky Jacky (280 metres a.s.].) and surrounded by coastal cliffs. Largely defoliated by feral rabbits, pigs and goats, the island is eroded extensively, with many exposed boul- ders and rock outcrops. The plateau above the coastal cliffs consists of a series of eroded valleys which rise towards the summit, where the slope ends abruptly at a near-vertical cliff, 280 metres a.s.l. (Coyne, 1982). Nepean Island, an uninhabited limestone island a few acres in extent, rises to 32 metres (Hoare, 1965). The climate of Norfolk Island is sub-tropical with no extremes of temperature and moderate relative humidity (72-81%) with no seasonal pattern. Mean monthly tem- perature fluctuates from minima near 12°C (in winter) to maxima near 25°C (in summer) and the daily range is rarely more than 8°C. Rainfall (mean annual precipita- tion 1313mm) occurs throughout the year with a winter maximum (June average of 163mm). Droughts of some weeks duration may occur (Ovington, 1984). HISTORY OF LICHENOLOGICAL INVESTIGATIONS Captain James Cook discovered Norfolk Island on October 10, 1774, during his second voyage around the world. The following day he landed at Duncombe Bay and named the island after the then Duchess of Norfolk. Although uninhabited at the time, Polynesian stone adzes have been discovered since, suggesting earlier settlement. Four- teen years after discovery, the first of two penal settlements was established and main- tained until 1814. A second penal settlement was established in 1826 and abandoned 30 years later. In 1856, descendants of the ‘Bounty’ mutineers left their home on Pitcairn Island and established the first free settlement on Norfolk, and today the ‘Islanders’ still comprise some 30% of the total population of approximately 2,000 (Hoare, 1965). Norfolk Island has been visited by a number of scientific expeditions in the course of its short history. Few of the early expeditions contributed greatly to the lichen flora. PROG. LINN. SOG. N.SW., 111 (2), 1989 J. A. ELIX AND H. STREIMANN 105 Captain Cook Mem. A Mt Bates NORFOLK ISLAND A Mt Pitt Cascade Ba 0} Flat Rock X Broken Pine y X "The Cockpit" Ball Bay Pt. Hunter 4 NEPEAN ISLAND PHILIP ISLAND 167°5S' Fig. 2. Map of the Norfolk Island complex. Here, and on Fig. 3, ‘Philip Island’ should be Phillip Island. In 1804-5 Ferdinand Bauer visited the island and his collections and drawings were submitted to Stephan Endlicher of Vienna, who published a Prodromus (Endlicher, 1833 to the plants which listed four lichens: Evernia (Usnea) melaxantha Ach., Parmelia (Parmo- trema) perlata Ach., Parmelia (Physcia) caesia Ach., and Sticta (Pseudocyphellaria) aurata Ach. In 1830 Allan Cunningham visited the island and augmented the knowledge of its PROG. LINN. SOG. N.S.W., 111 (2), 1989 106 LICHENS OF NORFOLK ISLAND 1 botany; his notes were published posthumously in Heward (1842). A lichen not enumer- ated by Endlicher was listed, namely Ramalina scopulorum Ach. Baron Ferdinand von Mueller subsequently investigated the botany of the island on the basis of specimens collected by Isaac Robinson, a resident and agent for the Sydney Botanic Gardens. The identity of the lichen collections was communicated subsequently to J. H. Maiden (see below). In 1904 J. H. Maiden (Director of the Botanic Gardens in Sydney) reported a total of 29 lichen species for Norfolk Island. This list was prepared by Edwin Cheel (Botanic Gardens, Sydney) after examining the collections of Maiden and Boorman (made during a visit to the island in 1902), and was supplemented by F. von Mueller (Mel- bourne), who supplied the names of six species identified by J. Muller of Aargau, Switzerland. This list is presented here using currently-accepted names for taxa (bold) followed, where appropriate, by the synonyms used by Maiden in brackets. [?] infers a dubious identification — according to Maiden. The species list included: Anaptychia ciliaris (L.) Korb. [?]; Catinaria grossa (Pers. ex Nyl.) Vainio {Patellaria grossa Mull. Arg.}; Catinaria versicolor (Fée) Sipman {Patellaria versicolor Fée}; Chiodecton perplexum Nyl.; Clathro- porina eminentior Nyl.; Coenogonium implexum Nyl.; Dirinaria confluens (Fr.) Awasthi {Physcia confluens Mitt.}; Glyphis verruculosa Zahlbr. {Glyphis verrucosa Knight}; Leptogium cyanescens (Rabenh.) Korb {L. tremelloides L.}; Letrouitia bifera (Ny1.) Hafelln. {Hetero- thecium biflorum Nyl.}; Ochrolechia pallescens (L.) Massal. {Lecanora pallescens Fr} [?]; Parmentaria ravenelli (Tuck.) Mull. Arg. {Parmentaria havenli Tuck.} [?]; Parmotrema chinense (Osbeck) Hale & Ahti [Parmelia perlatus L.]; Pertusaria sp.; Phyllopsora parvi- folia (Pers.) Mull. Arg. {Psora parvifolia Mull. Arg.}; Physcia caesia (Hoffm.) Furnr.; Physma byrsinum (Ach.) Mull. Arg.; Pseudocyphellaria aurata (Ach.) Vainio {Sticta aurata Ach.}; Pyrenula nitida (Weigel) Ach. [?]; Pyxine cocoes (Sw.) Nyl.; Ramalina farinacea (L.) Ach.; Ramalina glaucescens Krempelh. {R. levodea Nyl. var. fastigiata Mull. Arg.}. Ramalina siliquosa (Huds.) A.L. Sm. {R. scopulorum Ach.} |?]; Ramalina thrausta (Ach.) Nyl. {R. thrausta (Ach.) Fr.} [?]; Teloschistes flavicans (Sw.) Norm.; Usnea aurantiaco- atra (Jacq.) Bory {Evernia melaxantha Ach.} |?]; Usnea barbata (L.) Weber ex Wigg. [U. barbata Ach.}; Usnea florida (1L.) Weber ex Wigg. {U. barbata Ach. var. florida (L.) Fr.}; Usnea intercalaris Krempelh. Since then only scattered individual reports of lichens occurring in Norfolk Island have appeared in the literature, such as that of Usnea nexilis Motyka and U. propinqua Stirton (Motyka, 1938), Ramalina arabum (Dill. ex Ach.) Meyen & Flot. (Riedl, 1976) and from preliminary reports of the present work (Elix and Streimann, 1985). PRINCIPAL VEGETATION FORMATIONS The vegetation of the Norfolk Island complex was divided into five major for- mations: sub-tropical rainforest, open Avaucarta woodland, pastures and foreshores, weedy forests dominated by Pszdium and Olea, and Phillip Island (Elix and Streimann, 1985; Ovington, 1984). Brief notes are given below for these, together with the dominant lichen genera occurring in each formation. Sub-tropical Rainforest No doubt the Mt Pitt Reserve, with its remaining areas of sub-tropical rainforest, is the major habitat for lichens on the island and has the richest lichen flora. Dominant spermatophytes including Avaucaria heterophylla together with varying amounts of native hardwoods — Elaeodendron cirtipendulum, Nestegis apetala, Rapanea crassifolia and Baloghia lucida. ‘The understorey comprises smaller trees and sometimes scattered palms (Rhopalostylis bauert), tree ferns (Cyathea spp.) and the exotics red guava, Psidium Iittorale PROG. LINN. SOC. N.S.W., 111 (2), 1989 J. A. ELIX AND H. STREIMANN 107 and Citrus limon. Within this reserve, the eastern and south-eastern slopes of Mt Bates appear to be the richest site for lichens. Corticolous species abound: the richest sub- strates are the trunks of Elacodendron and the base of the trunks and branches of Araucaria heterophylla and Citrus limon. The macrolichen genera Heterodermia, Parmotrema, Ramalina, Teloschistes and Usnea are prominent in the canopy while Coccocarpia, Pannaria, Physma and Pseudocyphellaria species occur in the wetter areas at the base of trees. Elaeodendron curti- pendulum is by far the richest substrate for crustose lichens, including many species of Graphidaceae, Pyrenula and extensive cover by Clathroporina and Megalospora. One of the most conspicuous lichens of the forest is the beautiful Pseudocyphellaria aurata, with its brilliant yellow and emerald green (when wet) thallus, which commonly inhabits the trunks of Cyathea, Citrus and Araucaria. Open Araucaria Woodland Open Araucaria woodlands, which occur at sites such as Anson Bay Reserve, Selwyn Recreation Reserve and Ball Bay Reserve, constitute the second most important lichen habitat on the island. The Avaucarza trees throughout the island are characterized by a prodigious growth of Ramalina arabum and Usnea sp. on the canopy branches in par- ticular, as well as on lower branches. Dirinaria, Pyxine and Physcia species as well as various Ramalinae, are very abundant and well developed on the branches of most Araucaria trees, while Xanthoria and Caloplaca species are limited to Araucarza in coastal situations. Pastures and Foreshores Here old fence posts are colonized by a few corticolous lichens (Usnea, Ramalina, Lecanora sp.). Remnant Araucarza trees which occur in scattered stands or as individuals are much better substrates, with more species present and larger populations of lichens. However their communities are impoverished compared with those on woodland or forest trees of similar age. Exposed rock surfaces in pasture and along the foreshores are more interesting substrates. Their communities include representatives of Neofuscelza, Paraparmelia, Parmotrema and Xanthoparmelia plus a variety of microlichen genera includ- ing Diploschistes, Lecanora s.1. and Lecidea s.|. Interestingly the limestone outcrops near Emily Bay provide a suitable habitat for several calciphilic species, including a very unusual Buellia. Weedy Forest dominated by Psidium littorale and Olea africans Areas of Mt Pitt Reserve dominated by thickets of Pszdiwm littorale (guava) are extremely poor in lichens. Pszdium supports no lichens and its dense canopy prevents growth on soil or other small trees and shrubs. Lichens in these areas are found virtually only on scattered remnant Avaucaria or other native trees. Olea africans dominates other drier areas on the Reserve and, like Pszdium, it is a very poor lichen host. Only scattered depauperate Usnea, Ramalina and Parmotrema species are found on Olea in such areas. Phillip Island The Phillip Island lichen flora is rather depauperate because of the rarity of host trees. The branches of remnant Lagunaria patersonia are moderately well covered with lichens, but are relatively poor in taxa, mainly species of Dirinaria, Physcia and Pyxine. However saxicolous lichens are quite well developed in some areas of Phillip Island (e.g. upper Long Valley), although again the number of species is not large, with Parmotrema reticulatum being particularly common. Oddities include VTeloschistes flavicans growing on soil and rocks, usually a corticolous species, and Xanthoria, Caloplaca and Lecidea growing on consolidated soil, imitating the soil crust lichens of arid inland Australia. PROG. LINN. SOG. N.S.W., 111 (2), 1989 108 LICHENS OF NORFOLK ISLAND 1 THE LICHEN FAMILY PARMELIACEAE Representatives of the genera Flavoparmelia, Neofuscelia, Paraparmelia, Parmelia, Parmelinopsis, Parmotrema and Xanthoparmelia are detailed and a key to the species is given. A full description of each species (including chemical constituents) and its distribution are provided. Interestingly all (16) species have vegetative diaspores, illustrating the ease of dispersal of such units in reaching such isolated oceanic islands, and hence the effectiveness of such a mode of reproduction. Artificial Key to the Parmeliaceae in Norfolk Island cok Co ES OO ey pDihtalllhuisqlo nO wins iaectoe ht creed dane v at Mah pa ales cco Coe mea coy fe mea Neofuscelia verrucella dthallusseneysom yellow ence wes ms. er Ameen eer ai aorm ta ithe aren ea 2 ‘Bhallusiyellowsereent ss 42 epee RE a eat SA eds Ce ee ae 3 Mahala; eyes ee PEI RE TERIOR Iieiter Re RDN iar. AUR SERN Aya BAY es ORI eh aap OR de ea 5 sBhvallismvithraspaledloweigsuigialces i: ses aeesni eee ra Xanthoparmelia amplexula dBinallwsawatheat olackgloweryisuitclalce wry a eatee Oe rains ia eras itr ae yes a ae 4 Thallus with cylindrical isidia, medulla white (OV COMUCCA COMUNE RRA es tyas of place atarai nau chicane Git pradctcs aes gt Xanthoparmelia australasica Thallus with pustulate isidia, lower medulla WEllOW=Orralin oe tener iee sae OM Die nem TS ods sak, Be a Flavoparmelia norfolkensis ILOIOeS Marans ESS Waain I Bimnian ROC! 4 oy cellos C4 ob Ades bose s Webs eobae 6 ILOlnes loroacl, GReAior tnain Zisman lOPOAG! oo. 5 oo obs os Haddagideda Bou doe eo ale J divallusgisrdiatessaxicoloush ary ean heen el eee Faraparmelia scotophylla Thalllus joustullate, wsuallhy CormiColoOUs . 2505506050550 50005 Parmelinopsis spumosa Ujnoer swriace joseucloovplncllaite 2200 ¢a5 ccs scoues concourse va Farmelia erumpens Wippemsunacelackimegpscudocypiellacier ere ieee tetera re eae 8 Athallususidiateromsonediatea.2s ues +c oleate hha Maile a Wait meee eee 9 Thallus lacking soredia or isidia, but with flaking upper COOKE Was Ree eects Iaido tiirsbie nealing sais ade Parmotrema austrocetratum Pla ANI SHISTGII AWC teens, eset eee ute an stacey cout an UR Ae ivydtryin e (outa Cee gt ee eae ea 10 ablvalltisnsone diate mr tabs sees ie as occa tican eeuaret Ae ee et sek ar ene: eee eae 11 Winaillus culiaice, rmechuilley oe yell, (C= 4.05 co caccsesa0ccue Farmotrema crinitum Wells eeuliate, waecohuilley Ka (Cre Cl ao eo sca bo ooo soar soc Parmotrema tinctorum Wippermsunlacemencularelyacrackedinem em aera eae rene Parmotrema reticulatum Wippemsinracelackine retieulatereracks) sy em) sais einen ee eee te 2. IMieclWNAIC syalllow Or leas WEllOWReGl os eg oss acco nanesbsaoeacbseessors IS IVA TCSX I Baus) Se ig ay eas ir Sa ete rea eee Ly Htoe sR NSA Ae el 14 Medulla K+ yellow-red, containing salazinic acid ......... Parmotrema cristiferum Medulla K+ yellow, containing stictic acid ...........4.... Farmotrema chinense Medulla C+ rose, containing gyrophoric acid .......... Farmotrema sancti-angelit Mecha Crease ren). ae ena ey a ee oe a eee eee ie eo ee one 15 Thallus ciliate, medulla P-, UV+, containing alectoronic GIG EY ahs MoMA ihe Chee te ee shi Lo sad Team bea aes RE Farmotrema rampoddense Thallus eciliate, medulla P+ brick-red, UV-, containing protocetraric ACI. n ihe tigls Shs Hid MANET he Manny ini hae Sane CINE Ne ee eR oo Parmotrema gardneri PROG. LINN. SOG. N.S.W., 111 (2), 1989 J. A. ELIX AND H. SYREIMANN 109 Specimens Examined Collectors and location of specimens examined are as follows: JAE were collected by J. A. Elix and H. Streimann and are held in ANUC; HS were collected by H. Stretmann and are held in CBG with duplicates distributed as indicated. Collection site details (for map see Fig. 3): SIE open Araucaria woodland, Picnic Area, end of Martins Road, 29°03’30"S, 167°59’E, 80m, 1.x11.1984. 29°S NORFOLK ISLAND 4 NEPEAN ISLAND 29°05" 0 1 2km N @ PHILIP ISLAND 167°55S° 168°E Fig. 3. Collection sites on Norfolk Island. PROG. LINN. SOG. N.S.W., 111 (2), 1989 110 Swe S3: S4: S5s S6: SWE S8: S9: S10: Sil: S12: S13: S14: S15: S16: S17: S18: S19: S20: $21: S225 S85 $24: S25: LICHENS OF NORFOLK ISLAND 1 pasture, Cemetery road, Kingston, 29°03’30"S, 167°58’E, 15m, 2.xii.1984. Araucaria heterophylla-dominated grassland, Bloody Bridge, Cemetery Road, DONS HOGS, UOMO DS 18, Oren, Zs e4e, mixed exotic and Araucaria woodland, Rocky Point Reserve, 29°03’S, loo 20) 46 Ome Zax O82 mixed sub-tropical rainforest, near Broken Pine, Mt Pitt Reserve, 220-240m, DASA RO Sy MOP DO> ANE, Zoek, stand of Araucaria heterophylla along the margin of grassland and forest, Captain Cook Memorial, Duncombe Bay, 29°00'S, 167°56’30”E, 100m, 3.x11.1984. regrowth forest, Just south of the Captain Cook Memorial, Duncombe Bay, UOX00 20S, NOPD HO 18, UOOran, Src We. mixed sub-tropical rainforest, Filmy Fern Trail, Mt Pitt Reserve, 29°01'20”S, OPO 44018, Oro, Joan Oye, open woodland with kikuyu grass, Mt Pitt Reserve, track at end of Selwyn Pine IRGC ZOOS, O76" KOE, BilSvan, Sosa es, Olea africans-dominated valley, Upper Long Valley, Phillip Island, 29°07’30’S, SIDI Vd., BOiam, Fess". open Araucaria woodland, Point Blackbourne Reserve, end of Two Chimneys IRGaiel, 2°03 S, G79 1, Sian, Z.sou NM OSyee. mixed sub-tropical rainforest, Mt Bates summit trail, Mt Pitt Reserve, 29200 C3 0S 167,296) 3073 00m losxan O84 mixed sub-tropical rainforest, track from Red Road to Mt Bates, Mt Pitt Reserve, DOYO" AOS, IGT SG 440 1, P20 ma, Oasys. mixed sub-tropical rainforest, King Fern Valley, Mt Pitt Reserve, 29°01’S, NOP 2O 18, Aodhan, ovis, grassland with scattered Araucaria heterophylla, Flat Rock Bay Picnic Area, Anson Bay IRo@ael, ZO “SOS, IO 5H 18, Oi, Jasin OSH, Psidium- and Olea-infested lowland forest, track at end of Selwyn Pine Road, PIONS), MOP SG 308, 200mm, Bsr, rocky foreshore, Ball Bay Reserve, 29°03’S, 167°59’E, 2m, 8.x1i.1984. rocky cliffs at north-west point, Duncombe Bay, 29°00’S, 167°55’30”E, 50m, Oral O34. grassland with scattered Araucaria heterophylla, near Jacobs Rock, Anson Bay Re@aG!, LGC", IG 7° D518, S00, Osan ISSHe, Araucaria-dominated grassland, Bumboras Reserve, 29°03'30"S, 167°56’20”E, Oren, Osan iSe4e. open pasture, “(he Cockpit, Cascade Creek Valley, 29°01’S, 167°58°E, 35m, 9) SAMOS, open woodland, Prince Philip Drive, 29°01’S, 167°58’E, 35m, 9.xi1.1984. open pasture, Kingston, 29°03’30"S, 167°57'30"E, 8m, 9.x11.1984. regrowth rainforest, Mt Pitt Reserve, just south of summit of Mt Pitt, 29°01’S, 167°56’E, 230m, 10.x11.1984. mixed sub-tropical rainforest, Bird Rock ‘Track, Mt Pitt Reserve, 29°00'30"S, GJ D7 ld, AlOran, Osan Gey, FLAVOPARMELIA Hale Flavoparmelia norfolkensis Elix & Streim. sp. nov. (Fig. 4) Species cum thallo ut in Flavoparmelia euplecta sed ad hac specie isidiis cylindricis, coral- loideis demum inflatis, apicibus saepe erumpentibus sed esorediosis differt. Type: Norfolk Island. On Elaeodendron in open woodland in forest clearing, Mt Pitt PROC. LINN. SOG. N.S.W,, 111 (2), 1989 JA. ELIX AND. STREIMANN 11] INeschvcminackeatendrot Selwyn PimelRoad 29201 “Ssi167°d6 307 215mm, A. lax 18738 and H. Strermann, 8.xi1.1984 (CBG-holotype; MEL, US-isotypes). oy ng Pi we, pide So, x Fig. #4. Holotype of Flavoparmelia norfolkensis. Thallus foliose, corticolous or saxicolous, adnate, pale yellow-green, to 5cm in diameter. Lobes irregular, 1.0-3.0(-5.0)mm wide, becoming laterally imbricate, + contiguous, plane, rotund at the apices. Upper surface plane to + undulating, dull to shghtly shiny, with reticulate, white maculae towards the apices, isidiate; isidia laminal, cylindrical at first and becoming coralloid, ultimately becoming inflated at the apices and erumpent- pustulate, hollow within (dactylate), very rarely with traces of granular soredia; medulla white for the most part, but with a yellow-orange zone adjacent to the lower cortex. Lower surface black with a very narrow, naked, brown rim, sparsely rhizinate, rhizines short, simple or tufted at the apices, concolorous with the lower surface. Apothecia up to 5mm in diam., disc strongly concave, dark cinnamon brown, dull; margin persistent, strongly inrolled, pustulate-isidiate; spores ellipsoid, 19-21 x 8-94m. Pyenidia rare, black, punctiform, immersed; conidia weakly bifusiform, 4-6 x lum. Chemistry: Cortex K-; medulla K+ pale brown, C-, P+ brick red; pigmented lower medulla K+ claret; containing usnic acid, protocetraric acid (major), atranorin (+ trace), euplectin (minor), virensic acid (trace), gyrophoric acid (trace), skyrin (trace). PROG. LINN. SOG. N.S.W., 111 (2). 1989 112 LICHENS OF NORFOLK ISLAND 1 This species is characterized by the adnate, yellow to yellow-green thallus, the pustulate isidiate upper surface and a medulla pigmented orange (adjacent to the lower cortex) with the anthraquinone, euplectin. In many respects this new species resembles the common Australian species, / euplecta (Stirton) Hale, as both have a similar, pigmented lower cortex and analogous chemistry. However the upper surface of F euplecta is sorediate, with laminal, erumpent-pustulate soralia, which are subcapitate at first but coalesce and spread over the upper surface as granular soredia, whereas F norfolkensis 1s isidiate with the cylindrical isidia ultimately becoming coralloid, inflating at the apices and bursting open, but remaining esorediate for the most part. This species is quite common throughout the island. Specimens Examined: NORFOLK ISLAND. On tree trunk and dead log, $5, JAE 18333, 18352; on Elaeodendron, S7, JAE 18379; on canopy of fallen tree, $8, JAE 18421; on Elaeodendron, S9, JAE 18440, on Citrus limon, S12, JAE 18580; on fallen Araucaria heterophylla, S12, HS 34354; on dead treelet branch, S13, HS 34396; on Citrus limon, S14, JAE 18669; on volcanic rocks, S20, JAE 19269; on volcanic rocks, $21, JAE 18787, on Citrus limon, S24, JAE 18815; on rock shaded by old building, $23, HS 34806 (US). PHILLIP ISLAND. On volcanic rocks with a southerly aspect in the open, S10, JAE 16501. NEOFUSCELIA Esslinger Neofuscelia verrucella (Essl.) Essl., Mycotaxon 7: 53 (1978). Parmelza verrucella Essl., in C. F. Culberson, W. L. Culberson and T. L. Esslinger, Bryolo- ms S02 132 (O77). Type: Australia. Victoria: Rock Bore, about 33km NNW of Murrayville, Dahl (O-holotype). Thallus saxicolous, moderately to loosely adnate, yellow-brown to dark brown, 4-6cm in diam. Lobes irregular, 1.0-2.5mm wide, often markedly imbricate to entangled. Upper surface smooth, shining at margins, wrinkled and cracked in the centre, emacu- late, + lightly pruinose, moderately to densely isidiate; isidia laminal, cylindrical, sim- ple or branched, to 1.5mm tall; medulla white. Lower surface black, smooth and shining at margins, matt and wrinkled at centre, sparsely to moderately rhizinate, rhi- zines simple, black. Apothecia rare, to 2mm in diam., sessile, deeply concave when young, becoming + flat with age, disc smooth, light to dark-brown, margins entire or sparsely isidiate; spores broadly ellipsoid, 8-9 x 4-6 um. Pycnidia not seen. Chemistry: Cortex K-- HNO; + dark blue-green; medulla K-, C-, KC- or KC + faint rose, P-; containing divaricatic acid (major), nordivaricatic acid (minor). A common and widespread species in Australia, also occurring in both islands of New Zealand and South Africa. Rare in Norfolk Island. Specimen Examined: NORFOLK ISLAND. On volcanic rocks, S21, JAE 18791. PARAPARMELIA Elix & Johnston Paraparmelia scotophylla (Kurok.) Elix & Johnston, Mycotaxon 27: 281 (1986). Farmelia scotophylla Kurok., in Kurokawa and Filson, Bull. natn. Sci. Mus. Tokyo B, 1: 46 (1975). Type: On rocks, Ardglen Gap, Liverpool Range, 8.5km north of Murrurundi, Great Dividing Range, New South Wales, S. Kurokawa 5174, 28.x.1965 (TNS-holotype; MEL-isotype). PROG. LINN. SOG. N.S.W,, 111 (2), 1989 J. A. ELIX AND H. STREIMANN 113 Thallus saxicolous, adnate, mineral grey but blackening with age, 5-10 (-20)cm diam. Lobes irregular to sublinear, 1.0-3.0mm wide, slightly imbricate, apices subrotund. Upper surface smooth, shining at margins, flat to slightly convex, becoming cracked in the centre, emaculate, moderately to sparsely isidiate; isidia laminal, cylindrical, simple or sparingly branched, to 0.2mm tall; medulla white. Lower surface pale to light brown, sparsely to moderately rhizinate, rhizines simple, concolorous. Apothecia rare, to 6mm in diam., substipitate, deeply concave, disc smooth, pale brown, margins thin, involute, isidiate; spores 7-11 x 5-8 ym. Pycnidia not seen. Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange; containing atranorin, salazinic acid (major) and consalazinic acid. A common and widespread species in Australia, also occurring in both islands of New Zealand. Rare on Norfolk Island. Specimen Examined: NORFOLK ISLAND. On volcanic rocks, $21, JAE 18793. PARMELIA Acharius Parmelia erumpens Kurok., Lich. rar. Critic. Exsicc. no. 74 (1969) [Based on Parmelia tenuirama J. D. Hook. & Taylor f. corallina Mull. Arg.] Parmelia tenuirima J. D. Hook. & Taylor f. corallina Mull. Arg., Flora, Jena, 66: 46 (1883). Type: Gippsland, Australia, Starling (G-lectotype; UPS, US-isolectotypes). Thallus corticolous or saxicolous, adnate to loosely attached, pale greenish to light mineral-grey, 8-20cm in diam. Lobes short, subirregular to apically rotund, imbricate, 2-8mm wide. Upper surface shiny, plane, white-reticulate at first but becoming con- spicuously cracked to the margin, pseudocyphellae effigurate, 0.2-1.0mm long, some- what raised, dense, fusing into a reticulate network over the whole surface, sorediate; the soredia coarse and isidioid, often bursting apically, forming dense marginal and laminal soralia and/or extended, granular, coralloid-isidioid outgrowths; medulla white. Lower surface black, moderately rhizinate, rhizines simple or squarrosely branched at maturity, 1-2mm long, black. Apothecia rare, stipitate, to 15mm in diameter, disc dark brown, concave at first then flattening, margin inrolled at first but splitting radially at maturity, the amphithecium reticulately cracked, pseudocyphellate, sorediate; spores ellipsoid, 10-12 x 6-8um. Pycnidia scattered, punctiform, black, conidia cylindrical to weakly bifusiform, straight, 5-7 x lum. Chemistry: Cortex K+ yellow; medulla K+ yellow becoming deep red, C-, P+ red- orange; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (minor), lobaric acid (minor), protocetraric acid (+ trace). P. erumpens is a widespread species on rocks and trees in coastal and hinterland areas of Australia, both islands of New Zealand and Lord Howe Island. It also occurs in South Africa, India, Indonesia, Taiwan and Japan. This species is scattered on both rocks and trees in Norfolk Island and is easily recognized by the deeply reticulate- fissured cortex, the abundant coarse, pustular soredia and the lack of cilia. Uncommon in Norfolk Island. Specimens Examined: NORFOLK ISLAND. On exposed volcanic rocks, $2, JAE 16316, HS 31746; on Elaeoden- dron, 55, JAE 18310; on tree, S7, JAE 18360, 18385; on Elaeodendron, S16, JAE 18737. PARMELINOPSIS Elix & Hale Parmelinopsis spumosa (Asahina) Elix & Hale, Mycotaxon 29: 243 (1987). PROG. LINN. SOG. N.S.W., 111 (2), 1989 114 LICHENS OF NORFOLK ISLAND 1 Farmelina spumosa (Asahina) Hale, Phytologia 28: 483 (1974). Farmelia spumosa Asahina, J. Jap. Bot. 26: 259 (1951). Type: Higashi-Murayama, Kita-Iama-gun, Prov. Masashi, Japan, Asahina (1NS- lectotype). Thallus corticolous, adnate, fragile, pale mineral grey to pale olive grey, 2-6cm in diam. Lobes sublinear, narrow, 0.5-2.0mm wide, ciliate, the marginal cilia distinct and evenly dispersed, ca. 0.5mm long. Upper surface plane, emaculate, continuous, densely pustulate-isidiate; pustules laminal, bursting open but sparingly or not sorediate; medulla white or faintly yellow. Lower surface smooth, shiny, black, moderately rhizinate, rhizines simple or furcate, short, ca. 0.1-0.2mm long. Apothecia very rare, laminal, adnate, concave, 1-3mm in diam., disc brown, imperforate, margin and amphithecium pustulate; spores ellipsoid, 12-14 x 7-8um. Pyenidia not seen. Chemistry: Cortex K+ yellow; medulla K-, C+ rose, KC + red, P-; containing atra- norin, chloroatranorin, gyrophoric acid (major), 5-O-methylhiascic acid (minor), umbilicaric acid (minor). P. spumosa 1s a cosmopolitan species which is widespread throughout the tropical and sub-tropical areas of the world, but is much less common at temperate latitudes. In Australasia it is common in Australia, Papua New Guinea and New Zealand. Exposed trees at the forest margins or isolated individuals are the preferred habitat. Rare in Norfolk Island. Specimens Examined: NORFOLK ISLAND. On Citrus limon, S12, JAE 16584; on Elaeodendron, 322, JAE 18798; on Eiaeodendron, S25, JAE 18639, 10.x11.1984. PARMOTREMA Hale Parmotrema austrocetratum Elix & Johnston, Mycotaxon 31: 495 (1988) Type: New Zealand. North Island. On tree trunk in remnant forest, Burgess Park, New Plymouth, 60m, /. A. Elix 4645, 7v.1980 (CHR-holotype, CBG-1sotype). Thallus corticolous or saxicolous, loosely attached, light mineral-grey, 6-12cm in diam. Lobes rotund, imbricate or not, 10-20(-30)mm wide, apices often laciniate, the lacinae flat or convex, 0.5-1.5mm wide, 1-5(-8)mm long, lobes moderately ciliate, the cilia 0.2- 1.0(-1.5)mm long, simple or sparingly branched. Upper surface white-reticulate at first but becoming conspicuously cracked to the margin, developing raised closed dactyls, dactyls + curved, laminal, ultimately becoming fused and forming laminal ridges, the older parts of the thallus eventually becoming cracked-areolate, with the areolae flaking off and exposing the white medulla in eroded areas; lacking soredia and isidia. Lower surface black with a bare, brown marginal zone 1-2mm wide, moderately to densely rhizinate, rhizines simple or sparsely branched, slender, black. Apothecia rare, stipitate, to 10mm in diameter, disc perforate, pale tan, concave at first then concave- distorted, margin eciliate, thin, + stellate-cracked; spores ellipsoid, 12-16 x 8-9um. Pycnidia scattered, punctiform, black, conidia filiform, 9-16 x 1m. Chemistry: Cortex K+ yellow; medulla K+ yellow becoming deep red, C-, P+ red- orange; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (minor), protocetraric acid (+ trace). P. austrocetratum is a widespread species on rocks and trees in coastal and hinterland areas along the sub-tropical and tropical east coast of Australia and the north island of New Zealand. Uncommon in Norfolk Island. PROG. LINN. SOC. N.S.W., 111 (2), 1989 J. A. ELIX AND H. SUREIMANN 115 Specimens Examined: NORFOLK ISLAND. On Cyathea stem, 55, HS 31905; on Elaeodendron, 88, JAE 18439, HS 32190 (B, H, US); on Elaeodendron, S16, JAE 18742. Parmotrema chinense (Osbeck) Hale & Ahti, Taxon 35: 133 (1986). Lichen chinensis Osbeck, Dagb. Ostindisk resa,: 221 (1757). Type: Specimen and pl. 20, fig. 39B, Dillenius, Host. Musc.: 147 (1742) (OXF lectotypotype). Lichen perlatus Huds., Fl. angl.: 448 (1762), (incl. type of L. chinensis Osbeck). Farmelia perlata (Huds.) Ach., Meth. Lich.: 216 (1803). Parmotrema perlatum (Huds.) Choisy, Bull. mens. Soc. linn. Lyon 21: 174 (1952). Thallus corticolous, moderately to loosely adnate, membranaceous to coriaceous, pale mineral grey to whitish-grey, 4-15cm in diam. Lobes irregular, 3-8mm wide, in part rounded and deeply crenate, in part irregularly incised and laciniate, imbricate, sparingly to moderately ciliate, cilia 0.2-3.0mm long, + branched. Upper surface dull, smooth, sorediate; soralia submarginal, causing lobe margin to become revolute and suberect, ultimately appearing labriform, soredia granular; medulla white. Lower sur- face black, shining, with a broad, brown, naked marginal zone, moderately rhizinate, rhizines simple, to 2mm long. Apothecia very rare, laminal, substipitate, concave, to 7mm in diameter, disc pale brown to cinnamon brown, imperforate, margin thick, inrolled, sorediate; spores ellipsoid, 25-27 x 16-18um. Pyenidia not seen. Chemistry: Cortex K+ yellow; medulla K+ yellow, C-, P+ orange-red; containing atranorin, chloroatranorin, stictic acid (major), constictic acid (major), cryptostictic acid (trace), menegazziaic acid (trace), norstictic acid (trace). P. chinense 1s a cosmopolitan species which is widespread throughout the tropics and temperate areas. In Australasia it is common in Australia and New Zealand, but also occurs in Fiji and Papua New Guinea. It is very rare on Norfolk Island, and although it was the first ‘“Farmelia’ reported for the island (Endlicher, 1833), this may well have referred to P. reticulatum, a morphologically similar and common species. Specimen Examined: NORFOLK ISLAND. On treelet stem, S24, HS 34830 (US). Parmotrema crinitum (Ach.) Choisy, Bull. mens. Soc. linn. Lyon 21: 175 (1952). Parmelia crinita Ach., Syn. Lich.: 196 (1814). Type: North America, Muhlenberg (H-holotype). Thallus corticolous or saxicolous, coriaceous, adnate, pale mineral grey to grey-green, 5-6cm in diameter. Lobes subirregular, crenate or irregularly incised, imbricate, 2- 5mm wide, the margin ciliate, cilia simple or branched, 0.5-3.0mm long. Upper surface plane, emaculate, smooth to rugose with age, cortex fragile, isidiate; isidia laminal and marginal, short-cylindrical at first, + becoming coralloid, granulose or occasionally dissolving into soredia, often ciliate at the apices; medulla white. Lower surface black, with a narrow brown, naked marginal zone, densely rhizinate, rhizines slender, simple, to 1.0mm long. Apothecia and pycnidia not seen. Chemistry: Cortex K+ yellow; medulla K+ yellow, C-, P+ orange; containing atra- norin, chloroatranorin, stictic acid (major), constictic acid (minor), cryptostictic acid (trace), norstictic acid (trace), menegazziaic acid (+ trace), connorstictic acid (+ trace). P. crinitum is a cosmopolitan species, widespread in humid habitats in temperate and tropical regions. In Australasia this species 1s common in Australia, New Zealand and Papua New Guinea. Common in Norfolk Island. PROG. LINN. SOG. N.S.W., 111 (2), 1989 116 LICHENS OF NORFOLKISLAND 1 Specimens Examined: NORFOLK ISLAND. On base of Araucaria heterophylla, 51, JAE 18133; on base of Araucaria heterophylla, S4, JAE 16216, 16246, 16323, HS 31821 (US); on tree and dead branches, S5, JAE 18276, 16293; on Araucaria stem, S6, HS 32018 (US); on Elaeodendron S7, JAE 18584; on Lagunaria, $8, HS 32181 pr. p.; on Campsis grandiflora, S9, JAE 18469; on base of Araucaria heterophylla, Sil, JAE 18513; on Citrus limon, S12, JAE 18579, 18581, 18588, 18590; on treelet stem, S12, HS 34376; on palm and Citrus limon, S14, JAE 18670, 18679, 18692, HS 34515; on volcanic rocks, $17, JAE 16749; on volcanic rocks, $20, JAE 19268; on Citrus limon, S24, JAE 18812, 18817. Parmotrema cristiferum (Taylor) Hale, Phytologia 28: 335 (1974). Farmelia cristifera Yaylor, Hooker’s Lond. J. Bot. 6: 165 (1847). Type: Calcutta, Wallich (FH-lectotype; BM-isolectotypes). Thallus corticolous, adnate to loosely adnate, coriaceous, pale mineral grey, 3-10cm in diam. Lobes irregular, 5-20mm wide, rotund at the apices, entire or weakly crenate, imbricate or subascending at the margins, main lobes eciliate, lateral lobes and lobe axils not or sparingly ciliate, cilia 0.5-1.5mm long. Upper surface dull, emaculate, con- tinuous, sorediate; soralia mainly marginal on lateral lobes, sorediate lobes more or less ascending, soredia granular; medulla white. Lower surface smooth, shiny, black, with a broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, short, ca 0.1- 0.2mm long, coarse. Apothecia very rare, laminal, adnate, concave, to 3mm in diameter, disc brown, imperforate, margin thick; spores ellipsoid, 25-30 x 13-15ym. Pycnidia not seen. Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange-red; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (+ minor). P. cristiferum is a cosmopolitan species that is widespread throughout tropical and sub-tropical areas. In Australasia and Oceania it is common in Australia, Fiji, Papua New Guinea, Pitcairn Island, Raratonga and New Zealand. Common in Norfolk Island. Specimens Examined: NORFOLK ISLAND. On base of Avaucaria heterophylla, S1, JAE 18138; on dead branches and shrubs, $5, JAE 18279, 18282, 18301, HS 31908; on Elaeodendron stem, S7, HS 532061; on Elaeodendron, S8, JAE 18420, 18427; on Elaeodendron, S9, JAE 18444; on Citrus lumon, S12, JAE 18581, 18588, 18590, HS 34264 (B, H, US); on dead treelet stem, S13, HS 34397, 34414 (US); on palm, S14, JAE 18692; on treelet stem and stump, S14, HS 34515, 34516, on crown of Araucaria, S15, HS 34596; on Elaeodendron, S16, JAE 18735 (Lich. Australasict Exsicc. Fasc. 4: 88); on palm stem and dead tree, S16, HS 34633, 34674, 34675 (US); on Citrus limon, S24, JAE 18812. Parmotrema gardneri (Dodge) Sérusiaux, Bryologist 87: 5 (1984). Farmelia gardneri Dodge, Ann. Mo. bot. Gdn. 46: 179 (1959). Type: Brazil, Gardner (F H-holotype). Thallus corticolous, adnate to loosely adnate, coriaceous, pale mineral grey, 4-6cm in diam. Lobes irregular, 8-15mm wide, rotund at the apices, imbricate or subascending at the margins, crenate, irregularly incised or sublaciniate, eciliate or rarely sparingly ciliate, cilia 0.2-0.5mm long. Upper surface dull, emaculate, becoming rugose with age, continuous or + cracked, sorediate: soralia marginal, linear along the margins to sub- capitate on somewhat revolute lobes or short marginal laciniae, sometimes spreading submarginally, soredia granular; medulla white. Lower surface wrinkled, black, with a PROG. LINN. SOG. N.S.W., 111 (2), 1989 J. A. ELLX AND H. STREIMANN 117 broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, short, ca. 0.1- 0.2mm long, slender. Apothecia rare, laminal, adnate, concave, to 3mm in diameter, dise dark brown, imperforate, margin thick, eciliate, margin and amphithecium soredi- ate; sores ellipsoid, 18-22 x 8-10um. Pyenidia rare, punctiform, conidia sublageniform, 6-7 x lum. Chemistry: Cortex K+ yellow; medulla K+ pale brown, C-, P+ brick-red; containing atranorin, chloroatranorin, protocetraric acid (major), unknown fatty acids (+ minor). The presence of the marginal soralia and medullary protocetraric acid plus the absence of cilia distinguish this from other species of Farmotrema on the island. It is a pan- tropical species known from Africa, Australia, Papua New Guinea and South America. Rare in Norfolk Island. Specimen Examined: NORFOLK ISLAND. On Araucara trunk, S1, JAE 18156. Parmotrema rampoddense (Ny1.) Hale, Phytologia 28: 338 (1974). Parmelia rampoddensis Ny1., Acta Soc. Sci. Fenn. 26: 7 (1900). Type: Ramboda, Ceylon, Almquist (H-NY L 35555-holotype; S-isotype). Thallus corticolous, loosely adnate, coriaceous, pale grey to mineral grey, 10-20cm in diam. Lobes irregular, 5-20mm wide, rotund at the apices, crenate, ciliate, cilia con- spicuous, 3-6mm long, simple or bifurcate. Upper surface dull, emaculate, rugulose towards the centre, sorediate; soralia mainly marginal, linear, sometimes spreading submarginally, soredia farinose; medulla white, often pigmented orange-red adjacent to the lower cortex. Lower surface smooth, shiny, black, with a broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, to 6.0mm long, slender. Apothecia not seen in Norfolk Island material. Hale (1965) reports apothecia 3-10mm in diam., disc imperforate, amphithecium sorediate and spores 10-12 x 6-7um. Pyenidia not seen. Chemistry: Cortex K+ yellow; medulla K-, C-, KC + red, P-; pigmented lower medulla K+ purple; containing atranorin, chloroatranorin, alectoronic acid (major), a collatolic acid (major or minor), skyrin (+). P. rampoddense is a common and widespread pantropical species. In Australasia it is known from Australia and Papua New Guinea. Rare in Norfolk Island. Specimens Examined: NORFOLK ISLAND. On fallen branch; $5,/AE 16/25; on treelet stem, S14, HS 34532. Parmotrema reticulatum (Taylor) Choisy, Bull. mens. Soc. linn. Lyon 21: 175 (1952). Farmelia reticulata Vaylor, in Mackay, Fl. Hibern. 2: 148 (1836). Type: Ireland, County Kerry, near Dunkerron (Ff H-holotype). Thallus corticolous or saxicolous, loosely adnate, membranaceous to coriaceous, pale mineral grey to grey-green, 4-20cm in diam. Lobes irregular, 5-15mm wide, in part rounded and deeply crenate, in part irregularly incised and laciniate, imbricate or subascending at the margins, sparingly to moderately ciliate, cilia 0.2-3.0mm long. Upper surface dull, reticulately maculate and cracked, sorediate; soralia marginal, linear along the margins to subcapitate, commonly on short marginal laciniae, some- times submarginal and punctitorm, soredia granular; medulla white. Lower surface black, rhizinate or papillate to the margins or with a brown, naked marginal zone, densely rhizinate, rhizines simple or squarrose, to 2mm long, slender. Apothecia rare, submarginal to laminal, substipitate, concave, to 8mm in diameter, disc pale to mid- brown, imperforate or narrowly perforate, margin thick, eciliate, margin and amphi- thecium sorediate; spores ellipsoid, 13-18 x 8-11m. Pyenidia rare, punctiform, conidia filiform, 12-16 x 1.0-1.5um. PROC. LINN. SOG. N.S.W., 111 (2), 1989 118 LICHENS OF NORFOLK ISLAND 1 Chemistry: Cortex K+ yellow; medulla K+ yellow then dark red, C-, P+ orange-red; containing atranorin, chloroatranorin, salazinic acid (major), consalazinic acid (+ minor). P. reticulatum 1s a very common and highly variable species. Specimens growing in drier, exposed sites tend to become coriaceous, have weakly developed maculae and often submarginal soralia, while those from moist, shady habitats are usually membran- aceous, have marginal soralia and well-developed maculae. P. retzculatum is a cosmopoli- tan species, widespread throughout the tropics and temperate areas. In Australasia it is common in Australia, New Zealand and Papua New Guinea. Very common in Norfolk Island and occasional in Phillip Island. Specimens Examined: NORFOLK ISLAND. On Araucaria trunk, S1, HS 31716, 31723, 31728, 51734 (US); on exposed boulder, S2, HS 31748 (H, US); on small rock outcrop, $3, HS 51768, 31769 (B, H, US); on Grevillea robusta, and Melia, $4, JAE 18222, 16231, 18250, HS 31791, 31839 (US); on dead branch, $5, JAE 18308; on rocks, $6, JAE 18371, 18374, HS 31997, 32022 (US); on old wooden steps, $8, HS 32169 (H, US); on Cyathea stem, S14, JAE 16665, HS 34557 (US); on Araucaria trunk, S15, HS 34578A, 345868 (US); on volcanic rocks, S17, JAE 18745, 18746, 18837, HS 34695, 34699, 34707 (US); on volcanic rocks, S18, JAE 18764, HS 34736; on Lagunana patersonia trunk, S19, JAE 187768; on volcanic rocks, S20, JAE 16833 (Lich. Australasics Exsicc. Fasc. 4: 89), 186834, 19267, HS 34769 (1, US); on shaded rocks, S21, HS 34782. PHILLIP ISLAND. On south-facing volcanic rocks in the open, S10, JAE 18502, 18503, AS 32204. Parmotrema sancti-angelu (Lynge) Hale, Phytologia 28: 339 (1974). Parmelia sancti-angelu Lynge, Ark. Bot. 13: 35 (1914). Type: Santo Angelo, Rio Grande do Sul, Brazil, 25 January 1893, G. A. Malme (S-lectotype). Thallus corticolous, adnate to loosely adnate, membranaceous to coriaceous, pale grey to pale grey-green, 3-10cm in diam. Lobes irregular, 5-15 mm wide, rotund at the apices, crenate, often deeply divided and with ascending margins, ciliate, cilia slender, 1.0-3.5mm long, simple or bifurcate. Upper surface dull, emaculate, wrinkled or rugose towards the centre, sorediate; soralia mainly marginal, linear, sometimes spreading submarginally, soredia farinose; medulla white. Lower surface smooth, shiny, black, with a broad, brown or mottled, naked marginal zone, densely rhizinate, rhizines simple, elongate, to 2mm long, slender. Apothecia not seen in Norfolk Island material. Hale (1965) reports apothecia imperforate, spores 13-18 x 7-10um. Pyenidia rare, conidia weakly sublageniform, 6-8 x lum. Chemistry: Cortex K+ yellow; medulla K-, C+ pale red, P-; containing atranorin, chloroatranorin, gyrophoric acid (major), lecanoric acid (+ trace). P. sancti-angelit is acommon and widespread pantropical species. In Australasia it is known from Australia and Papua New Guinea. Rare in Norfolk Island. Specimen Examined: NORFOLK ISLAND. On Elaeodendron, 55, JAE 18349. Parmotrema tinctorum (Despr. ex Nyl.) Hale, Phytologia 28: 339 (1974). Farmelza tinctoria Despr. ex Nyl., Flora, Jena 55: 547 (1872). Type: Canary Islands, Despréeaux (H-NYL 35365-holotype). Thallus corticolous or saxicolous, loosely adnate, membranaceous to coriaceous, pale grey to grey-green, 10-30cm in diam. Lobes irregular, 10-20mm wide, rotund at the PROG. LINN. SOG. N.SW., 111 (2), 1989 J. A. ELIX AND H. STREIMANN 119 apices, entire or crenate, eciliate. Upper surface shiny, becoming dull towards the centre, emaculate, cortex sometimes cracking and flaking, isidiate; isidia sparse to abundant, laminal and eventually marginal, confluent or in scattered groups, brown tipped or concolorous with the thallus, simple or branched, thin and cylindrical or coarse and irregularly inflated, rarely + interspersed with lobules; medulla white. Lower surface smooth, shiny, black, with a broad, brown, naked marginal zone, sparsely rhizinate, rhizines simple, short, ca 0.5-2.0mm long, coarse. Apothecia very rare, laminal, substipitate, concave or + radially split, to 20mm in diameter, disc dark brown, imperforate or with a small perforation, margin thick, margin and amphithecium isidiate; spores ellipsoid, 13-15 x 7-8um. Pyenidia rare, conidia filiform, lGexle Ole opin: Chemistry: Cortex K+ yellow; medulla K-, C+ red, P-; containing atranorin, chloroatranorin, lecanoric acid (major), orsellinic acid (trace). Farmotrema tinctorum is a cosmopolitan species that is widespread throughout tropical and temperate regions. In Australasia and Oceania it 1s common in Australia, Fiji, Papua New Guinea, Vanuatu, Samoa and the North Island of New Zealand. Very common in Norfolk Island and Phillip Island. Specimens Examined: NORFOLK ISLAND. On base of Araucaria heterophylla, S1, JAE 18137, HS 31739 (H, US); on Grevillea robusta, S4, JAE 16221; on Elaeodendron and dead branches, $5, JAE 18285, 18307, 18350; on Araucania trunk, S6, HS 32017, 32055 (US); on canopy of fallen tree, S8, JAE 16411; on Elaeodendron, 88, HS 32164, 32196 (H, US); on Elaeodendron and Citrus limon, S12, JAE 16556, 18570; on Araucana trunk, S15, HS 34569 (B, MICH, US); on Elaeodendron, S16, JAE 18736 (Lich. Australasici Exsicc. Fasc. 4: 90); on rock outcrop, $20, HS 34660 (US); on tree trunk, $22, HS 36462 (B, US); on Citrus limon, 324, JAE 18616; on tree in open area, Mt Pitt Reserve, R. Goldsack 4, 25.x11.1981 (ANUC). PHILLIP ISLAND. On south-facing volcanic rocks in the open, S10, JAE 18499, HS 32225 (B, H, US); on Lagunaria stem, S10, HS 32213, 32225. VANTHOPARMELIA Hale Xanthoparmelia amplexula (Stirton) Elix & Johnston, Bull. Br. Mus. nat. Hist. (Bot.) 15: 192 (1986). Farmelia amplexula Stirton, Trans Proc. Roy. Soc. Victoria 17: 69 (1881). Type: Australia. Near Brisbane, Queensland, Bailey 262 (BM-holotype). Parmelia violascens Stirton, Trans Proc. New Zeal. Inst. 32: 77 (1899). Type: Australia. Grampian Mountains, Victoria, Sullivan (BM-holotype). Thallus foliose, adnate to moderately adnate on rocks, yellow-green or commonly blackening towards the centre as a result of the dark-tipped, dense isidia; subirregularly lobate, 5-10cm in diam. Lobes sublinear-elongate, sparingly imbricate or not so, 0.8-2.0 (3.0)mm wide. Upper surface opaque, emaculate, with sparse to numerous isidia, isidia cylindrical, simple or branched and coralloid up to 2mm high; medulla white. Lower surface pale brown to dark brown, sparsely to moderately rhizinate, the rhizines con- colorous with the lower surface, simple, slender. Apothecia rare, 1.5-6.0mm in diam.; disc strongly concave to + flat at maturity, dark brown, shining; margin thin, persis- tent, involute at first, isidiate; spores 7-12 x 5-6um. Pyenidia not seen. Chemistry: Cortex K-; medulla K-, C-, KC + rose, P-; containing usnic acid, loxodin (minor), norloboridone (major), unknown (trace). PROG. LINN. SOG. N.S.W., 111 (2), 1989 120 LICHENS OF NORFOLK ISLAND 1 A common and widespread species in Australia (occurring in all States and Terri- tories) and both islands of New Zealand. It also occurs in Lord Howe Island and South Africa. Rare in Norfolk Island. Specimen Examined: NORFOLK ISLAND. On treelet stem, $25, HS 349/64. Xanthoparmelia australasica D. Gall., N.Z. J. Bot. 18: 531 (1980). Parmelva australasica (D. Gall.) Filson, Aust. J. Bot. 30: 519 (1982). Type: New Zealand, North Auckland, Karekare Beach, on andesitic conglomerate, 8 October 1977, J. Bartlett (CHR 314047 — holotype; CHR — isotypes). Thallus foliose, moderately to loosely adnate, yellow-green, 5-12 (-20)cm in diam. Lobes irregular, 2.0-3.5 (-5.0)mm wide, often markedly imbricate, secondary lobes similar to the marginal lobes, sometimes building the thallus up into a thick mat, apices subrotund. Upper surface smooth, shining at margins, wrinkled and cracked in the centre, emaculate, moderately to densely isidiate; isidia laminal, often forming a dense areolate crust, robust, broad, cylindrical, extensively coralloid-branched, to 2.5mm tall; medulla white. Lower surface black with a narrow brown, naked, marginal zone, smooth and shining at margins, matt and wrinkled at centre, sparsely rhizinate, rhizines simple, black. Apothecia rare, 2-6 (-8)mm in diam., sessile, deeply concave when young, becoming shallowly convex with age, disc smooth, red-brown, margins con- spicuously isidiate, exciple wrinkled, shining, becoming isidiate with age; spores 8-11 x 4-6um. Pycnidia not seen. Chemistry: Cortex K-; medulla K+ yellow then dark red, C-, KC + red, P+ orange; containing usnic acid, salazinic acid (major) and consalazinic acid, protocetraric acid (+ trace), norstictic acid (+ trace). A common and widespread species in Australia, which occurs in all States and Territories. It also is found in both islands of New Zealand, the continental United States and South Africa. Relatively uncommon in Norfolk Island. Specimens Examined: NORFOLK ISLAND. On rock in shaded track cutting, S13, HS 543714; on volcanic rocks, S18, JAE 18765; on volcanic rocks, S21, JAE 18788, 18789, 18790, 18794, 18797, HS 34792 (US). ACKNOWLEDGEMENTS We thank the Australian National Parks and Wildlife Service for generous financial support for our field work on Norfolk Island, Dr G. A. Jenkins who determined the chemistry of many specimens and Ms Jen Johnston for many constructive suggestions to this manuscript. References Coyne, P. D., 1982. — Day of the rabbits. Studics on plant regeneration on Phillip Island. Geo 4: 30-39. ELIX, J. A., 1985. — Lichenes Australasici Exsiccatt. Fasc. 4. Canberra: Australian National University. , and STREIMANN, H., 1985. — Lichens and bryophytes of Norfolk Island. (Unpublished report to the Australian National Parks and Wildlife Service). ENDLIGHER, S., 1833. — Prodromus Florae Norfolkicae, sive Catalogus Stirpium quac in Insula Norfolk Annis 1804 et 1805 a Ferdinando Bauer collectae et depictae. Vienna: Fricdrich Beck. Har, M. E., 1965. — A monograph of Parmelia subgenus Amphigymnia. Contr. U.S. Natn. Herb. 36: 193-358. Hrwarb, R., 1842. — Biographical sketch of the late Allan Cunningham, Esq. Hooker’ Lond. J. Bot. 1: 107. Hoart, M., 1965. — Rambler’s Guide to Norfolk Island. Sydney: Pacific Publications. Jones, J. G., and McDouGALL, I., 1973. — Geological history of Norfolk and Phillip Islands, southwest Pacific Ocean. J. Geol. Soc. Aust. 20 (3), 239-257. PROG. LINN. SOG. N.S.W., 111 (2), 1989 J. A. ELIX AND H. STUREIMANN 121 MAIDEN, J. H., 1904. — Vhe flora of Norfolk Island. Proc. Linn. Soc. N.S.W. 28: 692-785. Moryka, J., 1936-38. — Lichenum gencris Usnea studium monographicum. I. Pars systematica. Leopoli (1.c. Lublin, Poland). OVINGTON, J. D., (director), 1984. — Plan of Management Norfolk Island National Park. Canberra: Government Printer, for Australian National Parks and Wildlife Service. RiEDL, H., 1976. — Beobachtungen an Ramalina arabum (Ach.) Meyen ct Flot. und Ramalinopsis mannii (luck.) Follm. ct Hun. Sydowra 28: 134-142. PROG. LINN. SOG. N.S.W., 111 (2), 1989 amine hei : apabieuabue part dalsiniiten sient od idle i “woth aaah: Fowwrdimys ANA since ian sic lehane ae banat? oc 1 we orev et ; DAT. thes : : a ee or Raalk tiga v5 heeeney o. yaiyh \ ipa carey: andl aM — PAL oy tidy 0 AF Pury ee ; Bipere.: sirtpli F.CesH) ae, 6h ipvigebs HIRATA ioe mally ec? 1 bia how pool ees inn wae any : Day uM Ave pin ue iyots ih Hi rath is 4K te et, at ce hora! etOyH ont: Da all sedeais APD nme (ying a y i ro j ily aisle ina We valid al ws - vic ah Sracrich Abd Ahecilike Spelt ree FU wo ‘v i ical perieby Cehaets - 4 tte + fe « Ny a re es i ihe ta): Bin) v a i : i Bd otecs 06 : 4 ra i a oo 1 fee Te te Mee a ee ee ee OO ee a visa eh han ona 7 reat ys, ee ee omen, Bante ge nes, ger Aw jis 1 cone A - ae ae Nts sss of Mae i ahs as ; nt ; ; ; fie =.) el hi ae ies E j eae § is > fi , 4 : 7 i A et ih 4 =) ie -) x 3 oy = eisai rad. ee) be peek lw: Ras ee eas uf rd ge tat rr ea ts ; ; ¥ os Taek ee ¢ : | ies AY ipeen ang NN eee Ber are a Aig Pia Fit ee. Te, Ae on, Ane ee ky) 7) > ¥ ri a = 1 ri y a ats. : 7 i), Fane Z ie ce a se » # ‘ we De Par. bea ti 4 a , M. 4 a : - ; Y ; : i : 4 A iia 7 rales ' Auditor Conicochernes doyleae, anew Australian Species of the Chernetidae (Pseudoscorpionida: Arachnida) CLARICE M. A. KENNEDY KENNEDY, C. M. A. Conicochernes doyleae, a new Australian species of the Chernetidac (Pseudoscorpionida: Arachnida). Proc. Linn. Soc. N.S.W. 111 (2), 1989: 123-129. A new species of the genus Conicochernes Beier, C. doyleae, is described from the Sydney region. It is corticolous, being associated in the sampled area specifically with the ‘Brush Box, Lophostemon confertus (R.Br.) Peter G. Wilson ct J. T. Waterhouse. Clarice M. A. Kennedy, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109, manuscript received 16 August 1986, accepted for publication 15 February 1989. INTRODUCTION The Chernetidae are the largest family of pseudoscorpions currently including nearly 100 genera and over 550 species distributed over much of the world (Harvey, 1985). Of these only 12 genera, represented by 26 species, occur in the Australian region. The species described herein is a representative of the genus Conicochernes Beier which is endemic to Australia. Four species are currently known namely: C. brevispinosus (L. Koch), C. crassus Beier, C. globosus Beier and C. incrassatus (Beier). Measurements are based on the examination of 10 specimens of each stage made in accordance with those advocated by Chamberlin (1931). Those in parentheses are the female and follow those of the male. Abbreviations for chelal trichobothria and setal formulae follow those devised by Chamberlin (1931). Genitalic terminology follows Legg (1974a, 1974b). All specimens are preserved in spirit. SYSTEMATIC DESCRIPTION CHERNETIDAE Chamberlin 1931 Conicochernes Beier 1948 Type species: Chelifer brevispinosus L. Koch 1885. For synonymy list refer to Harvey 1981: 246. KEY TOSPECIES OF CONIGCOCHERNES i orciistalibladcrottlacellumascrrates 26 34.52 ceed Ae Sele ss 2 Shekalistalgolac erotilacellmmypnotsenicatels aay eee) aes Cera 3 2, ENCICASE Z Galles SAAS, GS SOUS ORES =o 3 a se eee see geo C. doyleae sp. nov. icaleasctaMeyesspotsialseiter wei ay. erry a aeae sn tae: C. incrassatus (Beier) 3, omalahigtaliolacks ot ikereelikunes oyvarneitis 3. 455%s0ebe ues seen sssgeoneesssce-s 4 ANI ollevoles Ont ilavereliiluran jOUMIMAS . oes soc soe aasceaepsavccecae C. globosus Beier 4, Serrulla Sxiemor wile WOl7 lernellae 2. ese cesses: C. brevispinosus (LL. Koch) SEMAN with Zi Warne 2 so cache no see uscecenseoeass C. crassus Beier Conicochernes doyleae sp.nov. (Figs 1-4) Etymology: This species is named after Suzanne Doyle, a colleague who has been of great assistance to me in the field. PROG. LINN. SOG. N.S.W., 111 (2), 1989 124 NEW AUSTRALIAN PSEUDOSCORPION Fig. 1. Conicochernes doyleae sp.nov., scanning electron micrographs of paratypes: A, right chela, lateral aspect, male; B, left chela, lateral aspect, female; C, dorsal seta of tergite 4; D, serrula exterior of chelicerac, female; Da flagellum of left chelicera; E, movable finger of left chelicera. Holotype: 9, Australian Museum, Sydney (KS 17445). Type locality: Stanley Street, Chatswood, 10km NW of Sydney G.P.O., N.S.W., lat. 33°48°10"S, long. 151°11’30”E, under bark of Lophostemon confertus (R.Br.) Peter G. Wilson et J. T. Waterhouse, 17. iv. 1985, C. Kennedy. Paratypes: same data as holotype, Australian Museum, 40 (KS 17446), 49 (KS 19207), 4 nymphs (KS 19208). PROG. LINN. SOG. N.S.W., 111 (2), 1989 C.M. A. KENNEDY De Fig. 2. Conicochernes doyleae sp.nov., scanning clectron micrographs of female paratypes: F, trochanter of right pedipalp; G, carapace, dorsal aspect (pedipalps dissected); H, pleural membrane. PROG. LINN. SOG. N.S.W., 111 (2), 1989 126 NEW AUSTRALIAN PSEUDOSCORPION Diagnosis: A species of the genus Conicochernes with the following characters: trichobothrium zt widely separated distally from cst; serrula exterior with 16-17 equi- dimensionally broad, blunt lamellae and a long, narrow acuminate proximal lamella; 2 galea setae present, eye spots distinct. Description: Adults. Colour dark red-brown, body dorso-ventrally compressed, surface of carapace predominantly papillate; pleural membrane with closely set undulating longitudinal folds (Fig. 2H). Pedipalp stout, trochanter with gently rounded anterior margin, setae very long and acuminate proximally (Fig. 2F), elsewhere short, straight, longitudinally ribbed in upper distal third and terminating in 3-4 small denticles of different lengths (Fig. 1C); sculpture coarsely papillate; L/W ratio 1.15-1.40 (0°), 1.20- 1.40 (Q); femur stout, anterior margin relatively straight, posterior margin well rounded proximally, abruptly pedicellate, setae short, terminally denticulate as described above, 2.75-3.00 (0°), 2.56-3.00 (Q); tibia stout, anterior margin gently rounded, distinctly pedicellate 1.69-2.40 (oO), 1.63-2.33 (Q); chela very stout, chelal hand broad proximally, narrowing distally and longer than chelal fingers, lateral margins relatively straight, chela (with pedicel) 2.93-3.44 (0), 3.02-3.44 (9), chela (without pedicel) 2.43-3.17 (O°), 2.75-3.25 (Q ) x longer than broad. .Fixed chelal finger with 8 trichobothria (Fig. 4M, N), z¢ well separated distally from ist; movable finger with 4 trichobothria, venom tooth large; marginal teeth on each finger 40-50 contiguous, accessory teeth present; sense spots present; male chelal fingers with extensive medial gape (Fig. 1A); female chelal fingers without gape (Fig. 1B). Chelicera small with 5 setae, lamina exterior present, serrula exterior attached throughout with 18-19 (0°), 17- 18 (Q ) equal broad, blunt lamellae and a long acuminate proximal lamella (Fig. 1D); galea well developed, O with 3-5 small rami, 9 5-6 rami, in most cases 2 galea setae present (Fig. 1E); flagellum with 3 blades of unequal length, distal blade long with 8-9 small spines, approximately 3.40um in length, spaced 4.0-4.70um along the anterior face (Fig. 1Da). Carapace: anterior margin truncate; pars cephalica covered with trans- verse discontinuous ridges and corresponding furrows that may be modified by very small, transversely aligned papillae, the remaining part of carapace coarsely and densely papillate; posterior margin slightly angulate with centrally elevated disc, lateral margins slightly converging distally (Fig. 2G); setae short, terminally denticulate, 6: 10: 60-70 (O°), 6: 13-14: 50-60 (9 ); L/W ratio 1.05-1.16 (07), 1.00-1.16 (9) x longer than broad, eye spots present. Tergites: interscutal membrane in shallow folds, setae short, terminally denticulate; chaetotaxy: O& 12-14: 14:: 14-16: 14: 16-17: 16-18: 18-20; 18-20: 17- 18: 18-20: 14-16: ?; 9 10-14: 12-15: 16-17: 16-18: 18-20: 18-20: 18-20: 18-20: 18-20: 13-15: Qe @ ronseroaire, (Coxe cnaciotarays Cr 2; 53 24-25, Os Ss WOR2, Oz ae WD, Oz Ss WO=2, Os 3s NDAs © Ye Be WO=io, We Ze WB, ie Ze OQ. te Be Jos. We Ze IS, Malle genntalas close apodemes elongate, tapering; paired lateral apodemes extend ventrally and laterally to curve upwards terminally (Fig. 3K). Chaetotaxy: anterior operculum large, posterior margin with row of 7 acuminate setae curling into genital aperture, anterior to these, a row of 9 short setae above which are 3-4 long setae forming a pyramidal effect and surrounded by 11-12 very short setae; posterior operculum small with only 10-12 scattered setae (Fig. 31). Female genitalia: with lateral apodemes which provide for muscle attachment and support of the genital atrium; two median cribriform plates associated with the medium accessory glands and two lateral cribriform plates on the anterior wall of the lateral diverticula, each plate cuticular and perforated by pores; spermathecae two blind, digitate projections resulting from antero-dorsal evagination of median diverticulum. Scattered cells occur in epithelium of median diverticulum (Fig. 3L). Chaetotaxy: posterior margin of anterior operculum with row of 27-30 acuminate setae, long over small genital aperture, becoming short laterally, anterior to these and PROG. LINN. SOG. N.S.W., 111 (2), 1989 C.M. A. KENNEDY 27 0.1mm Fig. 3. Conicochernes doyleae sp.nov., 1 and J, scanning clectron micrographs of genital region, external aspect, male and female paratypes respectively. K, male genitalia; L, female genitalia. centrally situated 6-7 very long setae above which are 7-8 short setae; posterior oper- culum with 7-10 setae distributed centrally at random, posterior margin with 17-18 uniseriate acuminate setae (Fig. 3J). Sternites: setae short, acuminate; chaetotaxy: O (): 27-30: 12-14: 16-18: 20-23: 18-20: 18-20: 18: 18: 10-12: 10: °?; Q 0: 30-40: 20-24: 16-18: 16- SOG a loalO-18424-26) 282% stermites4-1l mmisermate: Dimensions (mm): body length 2.70-3.28 (2.92-3.46); pedipalps: trochanter 0.36- 0437 0227-0234 (0:36-0739/0:27-0:30), femur (027.7-0:90/0-27-0:30 (0:75-0584/0.27-0:30); tibia 0.59-0.72/0.30-0.39 (0.56-0.66/0.27-0.36), chela (with pedicel) 1.08-1.30/0.34-0.43 (1.18-1.26/0.36-0.41), chela (without pedicel) 1.02-1.24/0.34-0.43 (1.08-1.17/0.36-0.41), movable finger length 0.46-0.54 (0.43-0.56); chelicera 0.18-0.23/0.10-0.14 (0.21-0.25/0.11- 0.14) movable finger length 0.12-0.16 (0.12-0.18); carapace 0.84-0.98/0.75-0.89 (0.86- 1.03/0.84-0.98); cucullus length 0.14-0.19 (0.16-0.22) ocular breadth 0.56-0.64 (0.56- 0.64); leg I: coxa width 0.30-0.36 (0.32-0.36), trochanter 0.13-0.16/0.11-0.16 (0.14- 0.16/0.11-0.16), basifemur 0.16-0.22/0.14-0.18 (0.16-0.25/0.14-0.17), telofemur 0.22- 0.33/0.13-0.16 (0.22-0.33/0.12-0.16), tibia 0.16-0.28/0.10-0.14 (0.22-0.28/0.10-0.14), tarsus 0.16-0.22/0.08-0.10 (0.16-0.22/0.07-0.08); leg 4: coxa width 0.28-0.33 (0.30-0.36), trochanter 0.18-0.25/0.14-0.16 (0.19-0.28/0.14-0.19), basifemur 0.25-0.30/0.16-0.25 (0.19- 0.28/0.16-0.22), telotemur 0.39-0.50/0.19-0.24 (0.41-0.50/0.21-0.28), tibia 0.28-0.42/0.12- 0.14 (0.28-0.40/0.11-0.15), tarsus 0.22-0.28/0.08-0.10 (0.16-0.20/0.07-0.11). Tritonymph. L/W ratio: pedipalpal trochanter 1.36-1.73, femur 2.12-3.21, tibia 1.56- 2.05, chela (with pedicel) 3.00-3.65, chela (without pedicel) 2.78-3.30 x longer than broad. Fixed finger with 7 trichobothria, movable finger with 3 trichobothria, sb and ist absent; serrula exterior of chelicera with 13 broad lamellae, distal and proximal lamellae long and narrow. Carapace: posterior margin angulate, 6: 10: (50-60) setae, 1.04-1.20 x PROG. LINN. SOG. NSW... 111 (2). 1989 128 NEW AUSTRALIAN PSEUDOSCOR PION Fig. 4. Conicochernes doyleae sp.nov., female paratypes: M and N, scanning electron micrographs of right chela, lateral, external and internal aspects respectively showing position of trichobothria; P, Ist left leg: Q 4th left leg; R, left pedipalp, dorsal. PROG. LINN. SOG. N.S.W., 111 (2), 1989 C.M. A. KENNEDY 129 longer than broad. Tergal chaetotaxy: 10-12: 12: 12: 12: 13-14: 11-12: 14-15: 14-15: 12: 10- 12: 12-14: 2. Sternal chaetotaxy: 0: 6-8: 8-10: 12: 14-16: 16: 16-18: 10-14: 12-18: 10-12: 12- aC oxalehactotaxy. O2e44 Oi2 7/0 0250-7)) 02 wO=I3y Nlonotarsate: Dimensions (mm): body length 2.38-2.88; pedipalps: trochanter 0.26-0.33/0.19-0.22, femur 0.47-0.61/0.19-0.25, tibia 0.39-0.47/0.19-0.25, chela (with pedicel) 0.84-0.95/0.25- 0.30, chela (without pedicel) 0.78-0.86/0.25-0.30, movable finger length 0.35-0.45; carapace 0.61-0.70/0.56-0.64. Deutonymph. L/W ratio: pedipalpal trochanter 1.35-1.72, femur 2.14-2.57, tibia 1.56- 2.00, chela (with pedicel) 3.19-4.06, chela (without pedicel) 2.76-3.68 x longer than broad. Fixed finger with 6 trichobothria, movable finger with 2 trichobothria, st, sb, st, esb absent, serrula exterior of chelicera with 11 blunt lamellae, distal and proximal lamellae long and narrow. Carapace 6: 8: (30-40) setae, 1.00-1.07 x longer than broad. Wergal chaetotaxy: ‘8: 8: 8:-1012: 12: 10: 10-12: 12; 12: 10-112: 8: 2. Sternal chaetotaxy: 0: 0: 0: 4-6: 6-10: 7: 8-10: 9-11: 10-12: 9-10: 10: 1. Coxal chaetotaxy: 0: 2-3: 6-7, 0: 3: 5-6, 9227902937). Dimensions (mm): body length 1.48-2.24; pedipalps: trochanter 0.19-0.25/0.11-0.16, femur 0.30-0.42/0.14-0.19, tibia 0.25-0.33/0.12-0.19, chela (with pedicel) 0.61-0.73/0.16- 0.22, chela (without pedicel) 0.56-0.65/0.16-0.22, movable finger length 0.25-0.31; carapace 0).39-0.47/0.37-0.45. Protonymph. L/W ratio: pedipalpal trochanter 1.20-1.33, femur 1.75-2.50, tibia 1.58- 1.91, chela (with pedicel) 3.14-4.30, chela (without pedicel) 3.00-3.90 x longer than broad. Fixed finger with 3 trichobothria, ¢, et, eb, isb present. Serrula exterior of chelicera with 10 broad lamellae, distal and proximal lamellae long and narrow. Carapace with 6: 7-8: (25-30) setae, 1.04-1.10 x longer than broad. ‘Tergal chaetotaxy: 6: 8: 6: 6: 6: 8: 6-8: OCHO an otehnalachiactotaxyai0- 02-3214 40410-0110 0450-G410-6410-O510 1051-25 @.oxall Ghactoraxy 102410 geo O30 Ze 3. Dimensions (mm): body length 1.10-1.45; pedipalps: trochanter 0.12-0.16/0.10-0.13, femur 0.19-0.28/0.10-0.12, tibia 0.18-0.23/0.10-0.12, chela (with pedicel) 0.43-0.54/0.12- 0.14, chela (without pedicel) 0.39-0.48/0.12-0.14, movable finger length 0.18-0.21; carapace 0.28-0.48/0.26-0.43. ACKNOWLEDGEMENTS I wish to express my thanks to Dr P. D. Hillyard (British Museum (Natural History) ) for the loan of type material and to Dr M. Gray (Australian Museum) for reviewing the manuscript and offering comments thereon. References Brier, M., 1948. — Uber Pseudoscorpione der Australischen Region. Eos Madr. 24: 232-536. CHAMBERLIN, J. C., 1931. — The arachnid order Cheloncthida. Stanford Univ. Publy. (Biol) 7: 1-284. Harvey, M. S., 1981. — A checklist of the Australian Pscudoscorpionida. Bull. Br. arachnol. Soc. 5 (5): ZIP 2S ——,, 1985. — Pscudoscorpionida. Jn D. W. WALTON, (ed.), Zoological Catalogue of Australia, 3: 133. Canberra: Australian Government Publishing Service. LEGG, G., 1974a. — A gencralised account of the female genitalia and associated glands of pscudoscorpions (Arachnida). Bull. Br. arachnol. Soc. 3: 42-48. ——, 1974b. — A generalised account of the male genitalia and associated glands of pscudoscorpions (Arachnida). Bull. Br. arachnol. Soc. 3: 66-74. PROG. LINN. SOG. N.S.W,, 111 (2), 1989 oe ‘yh vit rie, i ee f wr ware ‘ a _flastiied | ! ‘ i Pic aay “ Wal te Mckenziartia and FPectocythere (Pectocytheridae, Ostracoda, Crustacea) in Lake Macquarie, New South Wales IRADJ YASSINI and MARY MIKULANDRA YASSINI, I., & MIKULANDRA, M. Mckenzzartia and Pectocythere (Pectocytheridac, Ostracoda, Crustacea) in Lake Macquarie, New South Wales. Proc. Linn. Soc N.S.W, 111 (2), 1989: 131-139) The distribution pattern of four species of Mckenziartia and one species of Pectocythere in Lake Macquaric, New South Wales, is investigated and three new species: Mckenziartia thomi sp. nov., Mckenziartia mowbrayi sp. nov. and Fectocythere royi sp. nov., arc described. Trad) Yassini, Dept of Geology, University of Wollongong, Wollongong, Australia 2500, and Mary Mikulandra, Dept of Geography, University of Sydney, Australia 2006; manuscript received 26 July 1988, accepted for publication 15 February 1989. INTRODUCTION The genus Mckenziartia was described by Bentley (1988) on the basis of carapace morphology. The genus was erected to designate a group of pectocytherid ostracodes characterized by the presence of a crescentic fossa over the central muscle field, a large anterior vestibulum with short, simple or occasionally-branched marginal pore canals, and an antimerodont hinge structure. The crescentic fossa over the central muscle field was described as a diagnostic feature of the genus. Examination of a large population of the species from different estuaries of New South Wales shows, however, that the size and shape of fossa may vary and is often integrated with the ornamentation pattern of the valves. The size and the shape of the vestibulum and the distribution of the anterior marginal pore canals remain unchanged, and they present a more stable diagnostic feature for the genus. Mckenziartia Bentley is a typical inhabitant of the estuarine environment and has been reported from several estuaries around Australia (Bentley, 1988). Present work investigated the distribution pattern of four species of Mckenzzartia: Mckenziartia portjacksonensis (McKenzie, 1967), Mckenziartia foveata (Hartmann, 1978), Mckenziartia thomi sp. nov. and Mckenziartia mowbrayt sp. nov., as well as distribution of Fectocythere royt sp. nov., another member of the Pectocytheridae in Lake Macquarie. The type specimens are deposited at the Australian Museum. Some paratype specimens were also deposited at the Commonwealth Palaeontological collection in the Bureau of Mineral Resources, Canberra. AREA STUDIED Lake Macquarie is located some 85km north of Sydney, on the central coast of New South Wales. With an extremely irregular outline, the lake extends in a north-south direction for over 22km, has a width of 10km and a surface area of 110km?. The average depth of the lake is 6.7m and its maximum depth never exceeds 12m. A narrow inlet channel 3.5km in length and 240m to 400m wide connects the lake to the Pacific Ocean. The lake has a total catchment area of 770km*. Three major creeks; Mannering Creek in the south, Dora Creek in the west and Cockle Creek in the north, contribute up to 4% of the total lake volume (Bass Becking et al., 1959). Some seven other minor creeks PROG. LINN. SOG. N.S.W., 111 (2), 1989 132 OSTRACODES FROM LAKE MACQUARIE pew CLD. pn Sik, ARGS Lake Macquarie Kel Built-up areas @ Sample location Vv Sample with Mckenziartia x No ostracodes Fig. I, Location map of the samples studied. PROC. LINN. SOC, N.S.W., 111 (2), 1989 I. YASSINIAND M. MIKULANDRA 133 TABLE 1 Species and number of valves in 30m of samples ; : Samples pipes BO | A ol SOP, Se VE ONE TT AE GE 8 Mckenziartia portjacksonensts 2... ... 26 10 245 Uae DES 8 ils} 40) 14 3 2 Mlekemznamtina WHORTD o555705000000¢ — — 48 = = = = pas aa = = = Mckenziartia foveata ............. _ — 12 — — = = = ae = = - Mckenziartia mowbrayt ........... = = D) =: a = ls 3 zs =e 8 [Pace MHORE THOME sooo adoacee doce. = — also flow into the lake during the rainy periods. A total of 26 grab samples were taken from the bottom sediment of the lake; Fig. 1 indicates the location of the samples studied. Table 1 shows the number of valves of each species in 30ml of unwashed sample. SYSTEMATICS Class OSTRACODA Latreille, 1802 Order PODOCOPIDA Muller, 1894 Family PECVOCYTHERIDAE Hanai, 1957 Genus Mckenziartia Bentley, 1988 MCKENZIARTIA PORTJACKSONENSIS (McKenzie, 1967) Fig. 2A-D and Fig. 4A ‘Hemicytheridea portjacksonensis McKenzie, 1967: p. 85, pl. 12, figs 31-j,6. Munseyella’ tumida Swanson, 1979: p. 158, fig. 4q. Fectocythere portjacksonensis (McKenzie), Hartmann, 1980: p. 122, pl. 5 fig. 17. FPectocythere portjacksonensis (McKenzie), Yassini and Jones, 1987: p. 826, pl. 2, fig. 21. Mckenziartia portjacksonensis (McKenzie), Bentley, 1988: p. 445, pl. 1, figs. e-f, text fig. 4a-b. Distribution: The species has been recorded from many estuaries and shallow open marine environments around Australia (McKenzie, 1967; Hartmann, 1978, 1980; Yassini and Jones, 1987; Bentley, 1988) and from New Zealand (Swanson, 1979). In Lake Macquarie, it was recorded from 12 stations but it was particularly abundant at station 47 at the mouth of the inlet channel (Table 1). Mckenziartia foveata (Hartmann, 1978) Fig. 2G-I, Fig. 3G and Fig. 4B FPectocythere foveata Hartmann, 1978: p. 144, pl. 14, figs. 12-13. Fectocythere foveata Hartmann, Yassini and Jones, 1987: p. 826, pl. 2, figs. 18-20. Remarks: The hinge structure and vestibulum pattern justify attribution to the genus Mckenziartia. Distribution: The species has been recorded from several estuaries and shallow open marine environments around the continent (Hartmann, 1978; Yassini and Jones, 1987). In Lake Macquarie it was only found in sample 47, dredged from a seagrass bed close to the inlet channel (Table 1). Mckenziartia thomi sp. nov. Fig. 3A-F and Fig. 4C Diagnosis: Carapace elongate, length about 2.6 times height. Anterior broadly rounded, posterior rounded ventrally, subvertical medio-dorsally. Dorsum straight, PROG. LINN. SOG. N.S.W., 111 (2), 1989 154 OSTRACODES FROM LAKE MACQUARIE Fig. 2. A-D, Mckenziartia portjacksonensis (McKenzic): A, RV, ©, external view; B, 6, LV, internal view; C, 9, ventral view of the carapace; D, 6, dorsal view of the carapace, sample 47. E, F, J-L, Mckenziartia mowbrayi sp. noy.; E, LV, 9, external vicw, holotype AM P37639; F, RV, 6, external view, paratype AM P37641; J, 9, ventral view of the carapace, paratype AM P37641; K, 9, dorsal view of the carapace, paratype AM P37642; L, LV, 6, internal view, paratype AM P37642, sample 1. G-I, Mckenziartia foveata (Hartmann); G, RV, 6, external view; H, 6, dorsal view of the carapace; I, LV, 6, internal view, sample 47. ventrum gently concave. Valves equal in size. In dorsal and ventral views, carapace elon- gate with parallel borders. Internally: inner lamella broad, anterior vestibulum deep, narrow, posterior vestibulum broad and elongate. Marginal pore canals simple, few and PROG. LINN. SOG. N.S.W., 111 (2), 1989 I. YASSINI AND M. MIKULANDRA 1335) Fig. 3. A-F, Mckenziartia thomi sp. nov.; A, LV, O,, external view paratype AM P37647; B, LV, 9, internal view, paratypc AM P37648; D, RV, 9, external view, holotype AM P37646; C, RV, oO’, internal view, paratype AM P37649; E, oO’, ventral view of the carapace, paratype AM P37650; F, o&, dorsal view of the carapace, para- type CPC 26554, sample 47. G, Mckenziartia foveata (Hartmann) ©’, ventral view of the carapace, sample 47. H-K, Pectocythere royi sp. nov.; H, RV, internal view, holotype AM P37661; I, LY, internal view, paratype AM P37662; J, dorsal view of the carapacc, paratype AM P37663; K, ventral view of the carapace, paratype AM P37664, sample 47. scattered. Hinge antimerodont. Muscle scars: four adductor scars in a subvertical row, two frontal scars and a large furcal scar. Valve surface reticulated, with three distinct PROC. LINN. SOG. N.S.W., 111 (2), 1989 136 OSTRACODES FROM LAKE MACQUARIE concentric ridges parallel to posterior margin and a set of oblique ridges joining the ventral ridge. Remarks: With a crenulate posterior socket, and a straight and crenulate median bar the species clearly belongs to the Pectocytheridae and it can be differentiated from the allied species (Mckenziartia portjacksonensis (McKenzie) and Mckenziartia mowbrayi sp. nov.) in the details of surface ornamentation. Material: Holotype AM P37646, right valve, adult male; length 0.51!mm. Paratypes AM P37647, AM P37648, AM P37649, AM P37650 and CPC 26554. All from type locality. Type locality: Lake Macquarie entrance tidal channel, New South Wales, Australia (lat. 33°03’00"S; long. 151°38’00”E); Recent, estuarine, muddy sand, with Zostera capricornt (Aschers.); depth 5.2m; on the sampling day salinity was 35.8°/o0, water temperature 22.8°C, and dissolved oxygen 6.6mg/I. Etymology: The species is named after Professor Bruce Thom, Geography Depart- ment, University of Sydney. Distribution: The species has also been recorded from Heron Island, off Queensland (Labutis, 1977) and Broken Bay, New South Wales (Yassini, unpublished). Mckenziartia mowbrayi sp. nov. Fig. 2E-F, J-L and Fig. 4D Diagnosis: carapace elongate, length about 2.3 times height. Anterior broadly rounded, posterior rounded to gently subacute. Dorsum straight, ventrum gently concave in the middle. Valves equal in size. In dorsal and ventral views, sides of the carapace are parallel. Internally: inner lamella broad, anterior vestibulum shallow and broad. Posterior vestibulum narrow. Marginal pore canals straight, well-spaced and few. Hinge antimerodont. Valve surface reticulate, with a few distinct, concentric low ridges at the posterior end, and three at the anterior end. In ventral view, the ridges form a series of chevron-type grooves converged toward anterior and posterior ends. Muscle scars are typical of the genus Mckenzzartia (Bentley, 1988). Remarks: When compared with Mckenzzartia thomi sp. nov, and Mckenziartia portjacksonen- sis (McKenzie), it differs in its typical surface reticulation. Material: Holotype AM P37639, carapace, male; length 0.46mm. Paratypes AM P37640, AM P37641, AM P37642, AM P37643 and CPC 26553. All from type locality. Type locality: Lake Macquarie (Sample 1), north of Mannering Park, New South Wales, (lat. 33°09'00"S; long. 151°33’00”E); Recent estuarine, sandy mud; depth 3.7m; on the sampling day salinity was 36.22°/o0, water temperature 22.8°C, and dissolved oxygen 4.4mg/1. Etymology: After Mr William Mowbray, Chairman of Lake Illawarra Management Committee. Distribution: Outside Lake Macquarie, the species has also been found in Broken Bay (Yassini, unpublished) and off Queensland (Labutis, 1977). PROC. LINN. SOG. N.S.W., 111 (2), 1989 I. YASSINIAND M. MIKULANDRA 13 100pm. Fig. 4. A, Mckenziartia portacksonensis (McKenzic), RV, internal structure; B, Mckenzvartia foveata (Hartmann), RV, internal structure; C, Mckenziartia thomi sp. nov., RV, internal structure, holotype AM P37546; D, Mckenztartia mowbrayi sp. nov., RV, internal structure, paratype AM P37644; E, Pectocythere royt sp. nov., LV, internal structure, paratype AM P37638. PROG, LINN. SOG. N.S.W., 111 (2), 1989 37 158 OSTRACODES FROM LAKE MACQUARIE Genus Pectocythere Hanai, 1957 Pectocythere royi sp. nov. Fig. 3H-K and Fig. 4E Fectocythere sp. (Ceduna 120), Hartmann; (Nec Hartmann, 1980) Yassini and Jones, 1987; Oe BAG, OL 2, ine, Weeclk/, Diagnosis: Carapace elongate, small, length about 2.1 times height. Anterior broadly rounded, posterior with an elevated rim and _ postero-ventrally oblique. Dorsum straight, slightly inclined posteriorly. Ventrum straight, valves ornamented with irregu- larly distributed rounded or elongated alveoh. In dorsal view, broader posteriorly. Inter- nally: inner lamella broad, anterior vestibulum broad and shallow, marginal pore canals few and straight. Hinge merodont, consisting in the right valve of an elongate anterior tooth, distinctly crenulated median groove and coarsely lobated posterior terminal tooth. Central muscle scars comprising a row of four adductors and a V-shaped antennal scar. Sex dimorphism is not known. Remarks: The only described species with closely similar shape is Pectocythere sp. (Ceduna 120) Hartmann (1980) which 1s easily distinguished from the present species by the depression of the postero-ventral region and lack of the raised posterior rim. Material: Holotype AM P37661, right valve: 0.43mm length. Paratype AM P37662, AM P37663, AM P37664 and CPC 26555. All from type locality. Type locality: Lake Macquarie (Sample 47), entrance tidal channel, New South Wales (lat. 33°03°00"S, long. 151°38’°00”E); Recent, estuarine, muddy sand with Zostera capricorni (Aschers.); depth 5.2m; on the sampling day, salinity was 35.8°/o0, tempera- ture 22.8°C and dissolved oxygen 6.6mg/I. Etymology: After Dr Peter Roy, Geological Survey, New South Wales. Distribution: Widely distributed in estuarine environments along the coast of New South Wales. DISCUSSION Among the five described species, Mckenzzartia portjacksonensis is the only one which is widespread and abundant in all the estuarine environments along the eastern coast of Australia; from Heron Island in the north (Labutis, 1977) to Bass Strait in the south (Yassini and Blom, in preparation). Pectocythere roy1, has also been found in limited numbers in Broken Bay, Twofold Bay and Bass Strait. Mckenzzartia foveata, initially described by Hartmann (1978) from Port Hedland, Western Australia, was also found in several estuaries of New South Wales (Lake Illawarra, Botany Bay, Broken Bay, Brisbane Waters). The species was also reported by Labutis (1977) from the Great Barrier Reef, off Queensland. The other two species; Mckenzzartia thomi sp. nov. and Mckenziartia mowbrayi sp. nov. appear to be limited to the northern (Labutis, 1977) and central coast of New South Wales. References Bass BECKING, L. G. M., THOMSON, J. M., and Woon, F. J., 1959. — Some aspects of the ecology of Lake Macquarie NSW with regard to alleged depletion of fish. I. General introduction. Aust. J. Mar, Fresh- water, Rey. 10; 297-305. BENTLEY, C., 1988. — Podocopida Ostracoda of Brisbanc Water, near Sydney, south-castern Australia. JT. HANAt, N. IKeYA and K. ISHIZAKI, (eds), Evolutionary Brology of Ostracoda: 434-448. Amsterdam: Elsevier. PROG. LINN. SOG. N.S.W., 111 (2), 1989 I. YASSINIAND M. MIKULANDRA 159 Hanal, T., 1957. — Studies on the Ostracoda from Japan. 1 — Subfamily Leptocytherinac n. subfam. /. Fac Sci. Univ. Tokyo Sect. 11 10: 469-82. HARtMANN, G., 1978. — Die Ostracoden der Ordnung Podocopida G. W. Muller, 1894 der tropisch- subtropischen Westkuste Australicns (zwischen Derby im Norden und Perth in Suden). Afitt, Ham Zool. Mus. Inst. 75: 53-219. ——., 1980. — Dic Ostracoden der Ordnung Podocopida G. W. Miller, 1894 der warmtempericrten und subtropischtropischen kustenabschnitte der Sud-und Sudostkuste Australiens (zwischen Ceduna im Westen und Lake Entrance im Osten). Mitt, Ham. Zool. Mus. Inst. 77: V1-204. Laputis, V. R., 1977. = Cytheraccan Ostracoda from the Great Barricr Reef. North Ryde, N.S.W.: Mac- quaric University, M.Sc. (Hons) thesis, unpubl. LArREILLE, P. A., 1802. — Histoire des Cypris et des Cytheres. In, LAPREILLE, P. A., (ed.), Histoire Naturelle Generale et Particulvere des Crustaces et des Insectes. Vol. 4: 232-254. Paris: Dulart. McKenzir, K., 1967. — Recent Ostracoda from Port Phillip Bay, Victoria. Proc Roy. Soc. Victoria 80: 61-106. MULLER, G. W., 1894. — Die Ostracoden des Golfes von Neapel und der angrenzenden Meeres-abschnitte. Naples Sta. Zool. Fauna Flora Golfes Neapel, Monographie, 31: 1-404. SWANSON, K. M., 1979. — Recent Ostracoda from Port Pegasus, Stewart Island, New Zealand. New Zealand J. Mar. Freshwater Research 13: 151-170. YASSINI, I., (in prep.). — Recent shallow water estuarine ostracods from New South Wales. ,and Bom, W., (in prep.). — Distribution of Recent ostracods in Bass Strait, south east Australia. , and Jones, B. G., 1987. — Ostracoda in Lake Illawarra, environmental factors, assemblages and systematics. Aust, J. Mar. Fresh. Research 38: 795-845. PROG. LINN. SOG. N.S.W., 111 (2). 1989 ve ie ia Ms ba Ate a is my ait Toll, pitta eae pisos Ree 3 Wo oie apd seaid ish slain Wa } bi inp ue ves fy clo ined oir lly é aye hk ER ia Sy Aha ea Ae Pp rey pecan) » awe ©: mn, satis rien e's apy? Leg nh eres el i < PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 111 NUMBER 3 Wetlands of the Lower Clarence Floodplain, Northern Coastal New South Wales IRS I, JPRIBSSIENZ (Communicated by P. ADAM) PRESSEY, R. L. Wetlands of the lower Clarence floodplain, northern coastal New South Wales. Proc. Linn. Soc. N.S.W. 111 (3), 1989: 143-155. The wetlands on the lower Clarence floodplain, on the far north coast of New South Wales, were mapped and described during a detailed survey in 1982. The survey identified 755 wetlands with a total area of about 14,700 ha. Most wetlands were smaller than 10 ha but the relatively few large wetlands made up most of the total area. Open water occupied only 5% of the total wetland area at the time of the survey although some open water occurred in 45% of wetlands. The most important plant families, in terms of extent and frequency of occurrence, were Poaceae (covering 23% of the wet- land area, occupying 82% of the total number of wetlands), Cyperaceae (20% area, 36% number), Myrtaceae (17% area, 36% number), Polygonaceae (12% area, 76% number) and Juncaceae (10% area, 75% number). Most wetlands were fringed by dense stands of herbaceous plants but, due to clearing, had few trees on their margins. Ninety-eight percent of wetlands were grazed to some extent and 92% of the total wet- land area was affected by drainage. Most wetlands are seasonal with relatively small catchments. Only 28% had catchments on the bedrock slopes surrounding the flood- plain. Most catchments have been completely cleared for agriculture. Most wetlands are freehold tenure. R. L. Pressey, New South Wales National Parks and Wildlife Service, PO. Box 1967, Hurstville, Australia 2220; manuscript received 1 November 1988, accepted for publication 15 March 1989. KEY WORDS: Wetlands, survey, plant taxa, catchments, Clarence floodplain. INTRODUCTION The coastal wetlands of New South Wales have been mapped and classified in a number of broad-scale surveys. Hannah’s (1968) landscape approach dealt mainly with the wetlands of the dune systems and was restricted to the north coast. West et al., (1985) mapped and classified the vegetation of the State’s estuarine wetlands. The surveys of Goodrick (1970) and the Coastal Council of New South Wales (1985) were of dunal, estu- arine and floodplain wetlands along the whole coast. The Coastal Council study simply delineated wetlands for the purposes of development controls under State Environmen- tal Planning Policy No. 14. Goodrick’s survey provided a useful classification based on dominant vegetation and water regime and is the best overview of wetlands on the coastal floodplains. It does not, however, provide sufficiently detailed information on individual floodplain wetlands for conservation or management. On the Clarence floodplain, none of the previous local, more detailed studies of flora and fauna has covered the wetlands comprehensively. They have concerned only one or a few wetlands and have had a variety of purposes and approaches. For example, Broome (1978) surveyed waterbirds in several areas and two environmental impact statements (Clarence River County Council, 1978, 1980) include vegetation maps and descriptions of some large wetlands. Most other publications on the Clarence floodplain wetlands deal with soils and specific occurrences of birds (Pressey, 1981). Wetland surveys for the National Parks and Wildlife Service, designed to be both detailed and comprehensive, covered the lower floodplains of the Hunter, Clarence and Macleay Rivers in 1981, 1982 and 1984, respectively, and the coastal lowlands of Tweed Shire in 1986. The main aims of these surveys were to: plot wetlands with accuracy and, in the PROC. LINN. SOC. N.S.W., 111 (3), 1989 144 LOWER CLARENCE WETLANDS Tweed survey, associated coastal communities on 1:25 000 topo-cadastral maps; describe the wetlands and their catchments from features that could be recorded during a single field visit or remotely from maps and aerial photographs; rank the wetlands for nature conservation, based on a combination of selected criteria. This paper summarizes the results of the inventory and description of wetlands on the lower Clarence floodplain. Full data, the results of conservation evaluation, and reduced wetland maps have all been incorporated in a consultancy report (Pressey, 1987). STUDY AREA The Clarence is one of the major coastal rivers in New South Wales and has an extensive floodplain, some 500 km north-east of Sydney (Fig. 1). The lower Clarence floodplain was defined as alluvium below the 10 m contour, a broad expanse of flood- prone land downstream of the very narrow, upper floodplain. Virtually all of the alluvial wetland area in the Clarence system occurs on the lower floodplain. The survey was also restricted to non-estuarine wetlands (those without saltmarsh and mangroves), although it included brackish areas intermediate between fresh and full tidal waters. The study area occupies approximately 700 sq. km (Fig. 1). To the north of the river, 1t extends upstream as far as Whiteman Creek and includes extensive alluvial flats to the north of Grafton, south of Lawrence and around the Broadwater. The major wet- land on this side of the river is the Everlasting Swamp. South of the river the survey covered areas downstream of Seelands including large expanses of floodplain between Swan Creek and Tyndale and around Wooloweyah Lagoon. The main wetlands in these areas are the Harrington Lagoon complex, the heavily wooded Shark Swamp and a group of large wetlands on the Coldstream River, one of the Clarence’s lower tributaries. The survey also covered Woodford, Harwood and Chatsworth Islands which lie either in the course of the river or in its broad estuary. Virtually all of the floodplain is cleared for farming. The major agricultural use 1s grazing of beef and dairy cattle. Sugar cane is an important crop in the eastern parts and relatively small areas are planted with poplars, vegetables and other crops. Drainage of wetlands began soon after European settlement. Around the turn of the century, drainage unions or trusts were formed to enable adjacent landholders to arrange for drainage in co-ordinated systems, with design and construction provided by the Depart- ment of Public Works. This work was to promote the growth of vegetation considered valuable as pasture and to encourage closer settlement in the region. From the late 1950s to the 1970s, under flood mitigation schemes, many drains, levees and floodgates were constructed and integrated to exclude floods from certain areas and to accelerate the recession of floodwaters. The alteration of wetlands continued during this period. Goodrick (1970) found that about 47% of all wetlands in the far north coast region had been destroyed or significantly altered by 1969, mainly by drainage and flood miti- gation. About 85% of the total affected area consisted of shallow floodplain wetlands, with more wetland now affected since flood mitigation on the Clarence has continued since Goodrick’s survey (Pressey and Middleton, 1982). In addition, many private developments have recently drained and cleared wetlands. METHODS The units of the survey were discrete wetland basins which were initially identified on black and white aerial photographs as areas of darker tone or as stands of trees. ‘The available aerial photographs were taken between August 1978 and March 1979 and varied in scale from 1:25 000 to 1:40 000. The minimum size for recognition of wetlands PROG. LINN. SOC. N.S.W., 111 (3), 1989 Re Ly PRESSEY 145 <— STUDY AREA Whiteman Creek 4 Coldstream River x. It SOUTH GRAFTON Fig. I. The study area. Numbers are those given to named wetlands during the survey. The names corres- ponding to these numbers are: 9 Double Swamp; 11 Bunyip Creek; 31 Alumy Creek; 54 Southgate Lagoon; 61 Southgate Creek; 62 Franks Creek; 72 Long Waterhole; 84 Hannons Lagoon; 90 Harrisons Creek/Coxs Creek; 123 Everlasting Swamp; 172 Mororo Creek; 181 Duckpond Creek; 206 Elbow Creek; 210 Roberts Creek; 257 Cowans Ponds; 278 Cassons Creek/Harrington Lagoon/McLachlan Waterhole; 302 Swan Creek; 306 Ellis Swamp/Crowsnest Swamp; 308 Horseshoe Waterhole; 400 Morans Swamp/Colletts Swamp; 413 Chaffin Swamp; 429 Champions Swamp; 436 Stokes Waterhole; 437 Stokes Waterhole; 464 Sweeneys Creek; 496 Sweeneys Swamp/Oregon Creek; 514 McPhees Swamp; 515 McPhees Swamp; 527 Calliope Creek; 573 Calligans Creek; 638 Alumny Creek; 649 Swampy Creek; 703 Shark Swamp/Tyndale Swamp. was about 0.1 ha. A few small wetlands of 1 ha or less were omitted because of difficult access. Final boundaries of wetlands were marked on aerial photographs during a single field visit to each one, either in summer (3.2.82-28.2.82) or winter (19.7.82-17.8.82). These boundaries were later transferred onto 1:25 000 topo-cadastral maps. Scale differences and lens distortion were corrected in the transfer by enlarging or reducing the images on the photographs and by matching features such as fencelines, roads and drains between photographs and maps over small areas. The number and types of attributes recorded for each wetland were constrained by the time available. In particular, fauna could not be described reliably, and faunal PROG. LINN. SOG. N.S.W,, 111 (3), 1989 146 LOWER CLARENCE WETLANDS habitat and importance in each wetland were generally inferred from open water and the type and structure of vegetation. Of the twelve attributes listed for each wetland, five were used solely for ranking the wetlands for conservation (see Pressey, 1987). The remaining seven were used descrip- tively and are as follows. 1) Szze Size of wetlands was estimated from corrected boundaries on topo-cadastral maps using a dot grid. i) Vegetation/habitats Vegetation and faunal habitat of the wetlands were described by plant species or genera and two categories of open water: deep (>60 cm) and shallow <60 cm). Authorities for plant names are those in Jacobs and Pickard (1981) as amended by Jacobs and Lapinpuro (1986), unless indicated otherwise. Water depth was judged from fence- lines and from the slope of the bed. Submerged plants were not recorded, nor were species from the families Azollaceae or Lemnaceae. The percentage of each wetland occupied by each taxon or open water category was estimated by eye in the field. Percen- tages were later converted to absolute areas when the sizes of wetlands were measured. ‘Taxa or open water categories occupying less than 0.1 ha in a wetland were listed only as present. Small wetlands were described by walking through or around them, large swamps by walking through them, by scanning with binoculars from vantage points, and by visiting areas with distinctive tone or texture on aerial photographs. Some inconsisten- cies in description would have resulted from differences in sampling intensity between wetlands. The accuracy of estimates of the percentages of wetland areas occupied by plant taxa and open water categories would probably be inversely related to wetland s1Ze. 11) Marginal vegetation Vegetation on wetland margins was rated in the field according to the percentage of perimeter lined with trees or emergents (rushes, tall herbs or grasses) and the average density and width of the fringing emergent band. Fringing plants not typical of wet- lands, such as tall pasture grasses, were not taken into account as emergents. iv) Alteration Signs of obvious alteration such as grazing, drainage and impoundment were recorded in the field and from aerial photographs. v) Catchment areas Wetland catchments were categorized according to whether they were completely on alluvial flats or at least partly on bedrock slopes. Catchments on bedrock above the 10 m (lowest) contour were delineated and their areas measured on topographic maps. v1) Catchment land use Broad types of land use in wetland catchments were recorded in the field and from maps and aerial photographs. They were listed for each catchment in order of the proportion occupied. Categories recognized were agriculture (including grazing and cropping), forestry, industrial areas, mining (only surface extraction of sand, gravel etc. in the case of the Clarence survey area), natural vegetation, urban areas and waste disposal (including sewage treatment works). vil) Ténure Land tenure, easements and any other relevant land designations were listed for each wetland in order of the proportion of the wetland affected. PROG. LINN. SOC. N.S.W., 111 (3), 1989 R. L. PRESSEY 147 RESULTS AND DISCUSSION The survey covered 755 wetlands with a total area of 14,728 ha. The majority of wetlands are small (Table 1), 88% of them 10 ha or less in size. These small wetlands account for only 9% of the total wetland area. The bulk of the wetland area (77%) is made up by the relatively small number of wetlands (3% of total number) greater than 100 ha in size. TABLE 1 Size distribution of wetlands No. of % Total Wetland % Votal Area (ha) wetlands no. area (ha) area 0-1.0 343 45.4 1749) 2 1.1-5.0 255 33.8 626.6 4.3 5.1-10.0 69 9,1 480.4 Boo 10.1-50 57 7.6 1155.4 7.8 51-100 12 1.6 878 6.0 101-500 13 tod 2713 18.4 501-1000 1 0.1 520 3.0) > 1000 5 0.7 8180 39).8) 755 100.0 14,7279) 100.0 Seventy-three plant taxa were systematically recorded. Their occurrence in the study area, with that of the two recorded categories of open water, is summarized in the Appendix. Some plants were identified only to genus. The most common of these were: — Juncus spp. refers to species in the Section Genuini, the relative areas of which were not determined. The only two species identified were /. usztatus and J. poly- anthemus. From subsequent work on the Macleay floodplain, it is likely that hybrids of these species are common and widespread. — Perswaria spp. were not recorded individually. Species collected during the survey were P. hydropiper (most extensive), P lapathifolia, P. strigosa and F. sp. B. Another species occurring in the area but not found in the wetlands surveyed is I& SO, A\ — Typha spp. were also combined. Both 7! domingensis and T: orientalis occur in the area, with the latter species by far the more common. Open water was relatively unimportant in terms of the overall area occupied (Appendix). Shallow open water was more extensive and more commonly recorded than deep open water. Goodrick (1970) recorded a relatively small area of wetlands domi- nated by open water on the far north coast generally. The floodplain wetlands in this region, including those of the Clarence, are largely in the five categories of Goodrick’s classification that are dominated by herbaceous emergents or by trees: fresh meadows, seasonal fresh swamps, semi-permanent fresh swamps, teatree swamps and reed swamps (‘Table 2). Wetlands in these categories contain relatively little open water. Of the plant taxa recorded, the monocotyledons accounted for nearly 59% of the total wetland area and dicotyledons about 35% (Appendix). Monocots occurred 1n 97 % of wetlands and dicots in 89%. The most extensive and commonly recorded monocot families were Poaceae (23% of wetland area, 82% of wetland number), Cyperaceae (20% area, 36% number) and Juncaceae (10% area, 75% number). Paspalum distichum (water couch) dominated the area occupied by the Poaceae and was the most commonly recorded grass. Eleocharis equisetina was the commonest and by far the most extensive member of the Cyperaceae. The most extensive and commonly recorded dicot families were Myrtaceae (17% of wetland area and 36% of wetland number) and Polygonaceae PROC. LINN. SOC. N.S.W., 111 (3), 1989 148 LOWER CLARENCE WETLANDS (12% area, 76% number). Melaleuca quinquenervia was the most extensive dicot species, occupying 17% of the total wetland area. Casuarina glauca was the most commonly recorded dicot species, occurring in 33% of wetlands, although it is likely that most or all of the 76% of wetlands containing Persicarza spp. contained FP. hydropiper. TABLE 2 Extent and decline of wetland types on the far north coast of New South Wales (modified from Goodrick, 1970)# Original 1969 To Wetland type area (ha) area (ha) Decline fresh meadows 22680 ( 2150) 59.4 seasonal fresh swamps ( 7050) semi-permanent fresh swamps 360 320 Hil. ih open fresh waters* 450 450 = teatree swamps 4010 2390 40.4 salt meadows 1380 1380 = reed swamps 1460 730 50.0 salt flats 530 530 = mangrove swamps 1860 1780 A) coastal bogs 570 570 = coastal Lepironia swamps 320 320 picdue 33620 17670 47.4 # estimates excluded two wetland types: shallow estuarine waters and sheoak swamps. * open fresh waters is the only floodplain wetland category identified by Goodrick as being dominantly open water, although relatively small areas of open water occur in other categories on the Clarence floodplain. Underlined categories are those occurring on the Clarence floodplain. Cyperaceae and Poaceae were the most diverse families, with 24 and 12 species respectively, together representing nearly half the total plant taxa. Most plant taxa occurred in relatively few wetlands and occupied only small percentages of the total wet- land area (Table 3). The maximum number of native plant taxa recorded in a wetland was 31, although most wetlands contained relatively few (Table 4). Four recorded plant species are introduced: Salvinia molesta, Echinochloa crus-gallt, Nymphaea capensis and Exchhornia crassipes (water hyacinth). At the time of the survey, these covered about 1.9% of the total wetland area. Water hyacinth was by far the most exten- sive of these, occupying some 250 ha, and had completely blanketed and infilled some wetlands. Several native species, such as Eucalyptus robusta, Juncus kraussi, Sporobolus virginicus and Trglochin striata, occur only marginally in floodplain wetlands and are more typical of other wetland types. The survey provided records of some rare or uncommon plant species. These include: — Brasenia schreberr (Gabombaceae): present in 1 wetland; very few records for coastal New South Wales; listed as 3V by Briggs and Leigh (1988); — Cyperus odoratus (Cyperaceae): present in 2 wetlands; very few records for the state; — Cyperus platystylis (Cyperaceae): present in 1 wetland; a plant of floating organic mats in northern Australia and a very rare plant in New South Wales (K. Wilson, Natnl. Herb. N.S.W., pers. comm.); found on a mat of water hyacinth; — Eleochars philippinensis (Cyperaceae): present in 1 wetland; only one previous record for the state; PROC. LINN. SOC. N.S.W., 111 (3), 1989 R. L. PRESSEY 149 TABLES Frequency of occurrence of plant taxa in wetlands according to Yo number of wetlands and % total wetland area (bracketed figures indicate number of introduced species in percentage classes) No. taxa No. taxa Percentage class (% no. wetlands) (% wetland area) 0-1.0 28(2) 57( 1.1-2.0 9 5(1) ZF) 5 2 3.1-4.0 1 2 4.1-5.0 8 1 5.1-6.0 Z = 6.1-7.0 2 — 7.1-8.0 2(1) — 8.1-9.0 = — 9.1-10.0 1 1 10.1-20 8 4 21-30 1 — 31-40 2(1) — 41-50 = = 51-60 = = 61-70 = _ 71-80 3 ~ 81-90 — _ 91-100 — = JUANBILIE, 2¢ Numbers of native plant taxa in wetlands Taxa No. wetlands % Total no. > 20 5 0.7 16-20 10 1 11-15 49 6.5 6-10 17 22.6 0-5 520 68.9 755 100.0 — Maunda triglochinoides (Juneaginaceae): present in 34 wetlands over a total area of 13.5 ha; known range only on the east coast between Wyong and the Brisbane area (Aston, 1973; Sainty and Jacobs, 1981); considered rare on the New South Wales central coast by Beadle et al. (1982); generally rarely reported and of uncertain status (S. Jacobs, Natnl. Herb. N.S.W., pers. comm.). Despite dry conditions over most of New South Wales in 1982, the results of the botanical survey should be generally representative of the Clarence floodplain wetlands. Good rains preceded the February survey when wetland vegetation was well developed and most basins were full or nearly so. However, some plants would have been missed or underestimated during the winter part of the survey because of seasonal growth patterns, absence of inflorescences, or the drier conditions typical of winter in the area. The majority of wetlands had few trees on their margins (Table 5), a result of the widespread clearing of the floodplain. Fringing herbaceous emergents were, however, generally well developed in terms of density, width and the average percentage of perimeters occupied. The major taxa classed as fringing emergents were Juncus spp. and Persicaria spp. Others were Carex appressa, Leersia hexandra and Lepironia articulata. Casuarina glauca and Melaleuca quinquenervia were the most common fringing trees. PROG. LINN. SOG. N.S.W., 111 (3), 1989 150 LOWER CLARENCE WETLANDS TABLES Summary of records for condition of marginal vegetation A B Percentage of perimeter with trees Percentage of perimeter with emergent vegetation Percentage No. wetlands % Votal no. Percentage No. wetlands % Votal no. 91-100 44 5.8 91-100 410 ee 66-90 36 4.8 66-90 V§) 10.5 36-65 58 70 36-65 74 9.8 10-35 65 8.6 10-35 42 5.6 0-9 552 73.1 ()-9 150 19.9 755 100.0 755 100.0 CC. Avcrage density and width of cmergent vegetation Rating No. wetlands % Votal No. 5 (very dense and wide) 331 43.8 4 69 9.1 3 122 16.2 2 76 10.1 1 (very sparse and narrow) 157 20.8 755 100.0 About 98% of the total number of wetlands were grazed to some extent. Drainage had directly affected 295 wetlands (39% of total number) and a wetland area of 13,500 ha (92% of total area). Drainage has reduced the persistence and depth of standing water in these wetlands with consequent changes in the distribution and abundance of plants and animals. The severity of drainage effects has varied depending on the depth of drains, the original nature of the wetlands, and the extent of their catchments. The overall effects of drainage on the Clarence floodplain wetlands are probably underesti- mated by these statistics for two reasons. Firstly, drainage may have lowered local water tables and so affected wetlands not directly drained. Secondly, drainage has fragmented some previously large wetlands into remnant sub-basins that are not directly drained but have had their water regimes altered. Few, if any, wetlands on the Clarence floodplain have been completely eliminated by drainage. The main reason for the inconsistency between this statement and Goodrick’s (1970) estimates of the decline of fresh meadows and seasonal fresh swamps in the region (Table 2) is that Goodrick’s survey was primarily concerned with waterfowl habitat. His estimates of the decline of wetlands therefore included any areas whose value to waterfowl had been largely eliminated. Many such areas, though, can still be defined by wetland vegetation. Wetlands were placed in two broad hydrological categories on the basis of the nature of their catchments. The first (Category 1) comprises 544 wetlands with an area of 2456 ha (72% total number, 17% total area) which have small catchments confined to alluvial flats below the 10 m contour. Their water levels would generally be less stable and would fall more quickly during dry periods than those in other wetlands. The second type of wetlands (Category 2) have catchments extending above the 10 m contour beyond the alluvial flats on which they occur. This category includes 211 wet- lands with a total area of some 12,272 ha (28% of total number, 83% of total area). Most of these wetlands have catchments smaller than 1000 ha although most of the wetland area in Category 2 is filled from larger catchments (Table 6). The largest catchment area, that of Ellis Swamp and Crowsnest Swamp (No. 306 in Fig. 1), 1s 17,200 ha. Most PROG. LINN. SOG. N.S.W,, 111 (3), 1989 R. L. PRESSEY 151 also have ratios of catchment area/wetland area of less than 100 and nearly 80% of the wetland area in Category 2 has a ratio of 10 or less (Table 7). Wetlands with large catch- ments and with large ratios of catchment area/wetland area will generally have more persistent and stable water levels and will contract more slowly in dry periods. At least some of these would be important local foci for fauna during droughts and may be significant in a broader context, particularly for the more mobile species of waterbirds. TABLE 6 Catchment sizes for wetlands with catchments above the 10 m contour (Category 2) Catchment No. % Total Wetland % Total size (ha) wetlands no. Cat. 2 area (ha) area Cat. 2 0-10 62 29.4 112.4 0.9 11-100 66 Sil 3} 421.3 3.4 101-1000 44 20.8 2566.8 20.9 > 1000 39 18.5 - 9171.4 74.7 211 100.0 12272.9 100.0 TABLE 7 Ratios of catchment area/wetland area for wetlands with catchments above the 10 m contour (Category 2) Catchment/ No. % Votal Wetland % Total wetland ratio wetlands no. Cat. 2 area area Cat. 2 0-10 106 50.2 9433.0 76.8 11-100 66 31.3 2722.8 22.2 101-1000 26 12.3 95.0 0.8 > 1000 13 6.2 21.1 0.2 211 100.0 122A 100.0 Twenty-one combinations of land uses were recognized in wetland catchments. The occurrence of each broad land use is summarized in Table 8. Agriculture is by far the major catchment land use, although areas of natural vegetation are also relatively common. Seventy-seven percent of catchments are completely under agriculture, and nearly all of these are restricted entirely to the alluvial flats. Only two catchments are completely covered by natural vegetation. TABLE 8 Land use occurrence in wetland catchments Present (% no. Dominant or sole Land use catchments) use (% no. catchments) Agriculture 99.7 5) Forestry 5.7 Holl Industrial area 0.5 Mining 357 = Natural vegetation Zl) UGS) Urban area 3.0 0.1 Waste disposal 0.5 = In 95% of wetlands, land tenure is solely or dominantly freehold. One wetland (Cowans Ponds, No. 257 in Fig. 1) is within a Wildlife Refuge under the National Parks and Wildlife Act, 1974. PROC. LINN. SOC. N.S.W., 111 (3), 1989 152 LOWER CLARENCE WETLANDS The survey reported here is the first comprehensive and detailed description of wet- lands on the Clarence floodplain. However, the results are accurate for the wetlands and their catchments only for the period of the field survey and for the dates of the aerial photography. Subsequent developments and changes in land use will have affected and will continue to affect the condition of many wetland catchments, the condition of marg- inal vegetation, and the occurrence of plant taxa and open water. In particular, the future applicability of the data on plants and open water categories will depend on the extent of short-term irregular and long-term successional changes in the wetlands. The nature of these changes ts difficult to predict with the presently poor understanding of the dynamics of Australian wetlands. Nevertheless, general observations by the author over several years on the Clarence floodplain suggest that: — the distribution and abundance of plant taxa in some wetlands alter over periods of months or years, although the suites of plant species in these areas generally remain similar; — the distribution and abundance of plant taxa in other wetlands, particularly those which are seasonal and have small catchments, appear to change very little over months or years. Without research on the vegetation dynamics of these areas, the characteristics of the wetlands that determine variability or stability of plant distributions, and the nature and extent of short-term variability, must remain conjectural. ACKNOWLEDGEMENTS The study was funded by the N.S.W. National Parks and Wildlife Service as a consultancy. Phillippa Pressey provided much valuable assistance during compilation and checking of the survey data and typed the original data base. Chris Ann Urquhart assisted in the preparation of the tables and drafted the figure. John Porter organized the analysis of the numerical distribution of open water categories and groups of plant taxa given in the Appendix. The staff of the National Herbarium of New South Wales, particularly Surrey Jacobs and Karen Wilson, assisted with the identification of plant specimens and with advice on their taxonomy and ecology. Surrey Jacobs, Karen Wilson and an anonymous referee provided helpful comments on the manuscript. References ASTON, H. I., 1973. — Aquatic Plants of Australia. Melbourne: Melbourne Univ. Press. BEADLE, N. C. W., EvANS, O. D. and CAROLIN, R. C., 1982. — Flora of the Sydney Region. 3rd edition. Sydney: Reed. BRIGGS, J. D. and LEIGH, J. H., 1988. — Rare or threatened Australian Plants — 1988 Revised Edition. Aust. Nat. Pks. Wildl. Serv. Spec. Publ. 14. BROOME, L. S., 1978. — Birds on north coast wetlands. Univ. New England Sch. Nat. Res. Rep. PRY. CLARENCE RIVER COUNTY COUNCIL, 1978. — Sportsmans Creek — Everlasting Swamp Environmental Impact Statement. CRCC: Grafton. ——.,, 1980. — Upper Coldstream Area Environmental Impact Statement. CRCC: Grafton. COASTAL COUNCIL OF NEW SOUTH WALES, 1985. — Coastal Wetlands of New South Wales. Department of Environment and Planning: Sydney. GOoDRICK, G. N., 1970. — A survey of wetlands of coastal New South Wales. CSTRO Div. Wildl. Res. Techn. Memo. 5. HANNAH, B. C., 1968. — The landscape of the north coast of New South Wales. Univ. New England Dept. Geogr. Res. Series in Appl. Geogr. No. 14. Jacoss, S. W. L. and LAPINPURO, L., 1986. — Alterations to the census of New South Wales plants. Zélopea 2: 705-714. , and PICKARD, J., 1981. — Plants of New South Wales: A Census of the Cycads, Conifers and Angiosperms. Government Printer: Sydney. PRESSEY, R. L., 1981. — A Review of Literature on the Floodplain Wetlands of Coastal New South Wales. National Parks and Wildlife Service: Sydney. PROC. LINN. SOC. N.S.W., 111 (3), 1989 Reis PRESSEN 153 ——, 1987. — A Survey of Wetlands on the Lower Clarence Floodplain, New South Wales. National Parks and Wildlife Service: Sydney. , and MIDDLETON, M. J., 1982. — Impacts of flood mitigation works on coastal wetlands in New South Wales. Wetlands (Australia), 2: 27-44. SAINTY, G. R. and Jacoss, S. W. L., 1981. — Waterplants of New South Wales. Sydney: Water Resources Commission. WEST, R. J., THOROGOOD, C. A., WALFORD, T. R. and WILLIAMS, R. J., 1985. — An estuarine inventory for New South Wales. N.S.W. Dept. Agric. Fisheries Bull. 2. PROG. LINN. SOG. N.S.W., 111 (3), 1989 LOWER CLARENCE WETLANDS 154 su oe 0'¢ 6% unyofin) unjpcygorucy Cr BITS 9°41 Orr punasinba -q AVAOVOVAOTVH a 2 v9 8h REDE D SOLAIOY GL It 8° 109 1s GZ ponvya DuisonsDy) ee ‘ Ness ; ee : AVAOVNIYVASVO ye © fa 6% Le SUR PASE, *O) su + 0) i sycsqnd “2D su + a0) I LagaLyIs DIUasDLT et 6-0 a I snsond ‘5 AVAOVANOEVO su I ©) % snjDvLopo “FD 10 COT al 6 p1j0f1douoL09 vjnjory su + £0 C uvodsvy “+ AVAOVUAALSV su VG vv ones snyoyjoxa snsagay) su + L‘0 C pyjuntog sisqoanjry ae PG 20 v winsaroug winipvi) AVAOVIdV Z0 9°46 69 ZS OSSAGAD AEE) SIU 8¢ 8G IG syuvranyt g SNOGATALOOIC Z'0 Z'GZ £0 z Myjampyp9 § snuaoyasoqog = = = SW pa Kjos OUST TEI6 su + 60 L DIDINIIYAD vaUNDE 9°89 1°£698 1°26 €€Z Pel W [23°.L AVAOVUAIAD 60) FG BE 66 ‘dds vydd7, su + 0 iG SnuIUu wniUosDULDG AVAOVHdAL AVAOVLVWNSITV su 9 I+ 1¢ SNJOULUD IA) UOJaTOULD}OT SNOCGATALODONOW AVAOVNO.LYUDOWV.LOd ; : 10 rst LG 6F sayAydopriayg [20], Ly L9%G GSE 99¢ Saq1SSD1I DIULOYYNGT AVAOVIaaCdaLNOd su G 40 6 IPA “S °C 62S 48 GHOSE =— HSB 229 avaoeod [210.1 eae et panies ues =e = : AVAOVINIATVS 0) tI 0 v snoqursara snjoqosogy 10 ETI I? 1¢ “VIA VIINU DapsLDI su + €'0 Z poipur sidajo100¢ AVAOVATISUVIN es CGY ly? Ig SUPE SUD Sel su + Oy OL ‘wang wnsipur wnuysag 7 Lape I'd 8 unjoursDa ‘q AVAOVNHOA TE 0ST CLIC’ bGL 696 wunysipsip wnjogsog Sa LAHAOGINALd 1Z + 60E 102 GGT Stud Dsopnasq/uimaruyg () su + €0 Z unsopnjng wingUDdg 0°S 028 O'FF L&E 12}@M uado [230], G0 C UG VL 9G YER] EDT 1G Cole erat 16 Jovem undo dooq Ste G0 (om 6 pprydojvujay “y 8 & CG vGG 0° GF LIE toyem uodo Morfeys Role potdnsa90 ‘ou spurp}oM ALOB9IVI BOIe potdna90 ‘Ou spuep1omM Ar0$91e9 purpom (ey) ese [Roy % ‘ON 19)}@M UddO/UOXe J, purpom (ey)eoie Roy % ‘ON 19}e8M uddo/uOXe J P10. % [210.1 [RIOL % [QUE (YT uoy? sap sayvoipur su “vy ['() Uvy) Lajvade aguasingzo ou savour + 1000 3 0.7 9370 73.4 432 100.0 12771.8 100.0 Open water was relatively unimportant in terms of the overall wetland area occupied (Appendix). Some shallow open water occurred in about half the number of wetlands but totalled, at the time of the survey, only about 290 ha or 2.3% of total wet- land area. Deep open water was much more restricted in distribution and extent. The scarcity of open water in the wetlands of the mid-north coast generally is indicated by estimates from Goodrick’s (1970) report (Table 2). The five categories of Goodrick’s classification that make up the bulk of the wetland area in the region, and on the Macleay floodplain, contain relatively little open water and are dominated by her- baceous emergents and by trees. Completion of all the field work in mid-summer following heavy rains provided ideal conditions for the identification of a large number of wetland plant taxa. Some shallow seasonal wetlands contained so much water that the results of the survey may be atypical to some extent. Of the plant taxa recorded, one 1s a liverwort, seven are ferns, 67 are monocotyledons and 43 are dicotyledons. The most diverse families were Cyper- aceae (30 taxa), Polygonaceae (12), Juncaceae (11) and Poaceae (10). The largest number of native taxa recorded in a wetland was 46, although most wetlands contained relatively few (Table 3). There is a significant relationship between wetland size and number of plant taxa in the wetlands on the Macleay floodplain (Margules et al., 1988) although there is considerable variability in the number of taxa in wetlands of any size. Most taxa occurred in relatively few wetlands and occupied a relatively small overall area (Table 4). Monocotyledonous plants occupied some 62% of the total wetland area and occurred in all but one wetland (Appendix). The most important monocot families, in terms of extent and frequency of occurrence, were Poaceae (28% of total wetland area, 91% of total number of wetlands), Juncaceae (10% area, 91% number) and Cyperaceae (22% area, 63% number). Paspalum distichum (water couch) and Juncus polyanthemus x usitatus were the most extensive and frequently occurring taxa in the Poaceae and PROG. LINN. SOG. N.S.W., 111 (3), 1989 R. L. PRESSEY 161 _Juncaceae, respectively. Eleocharis equisetina made up most of the area occupied by the Cyperaceae and was one of the most frequently occurring sedges. Dicotyledons occupied 35% of the wetland area and occurred in all but three wetlands. Polygonaceae (26% area, 97% number) and Myrtaceae (6% area, 21% number) were the major dicot families. Persicarta hydropiper and Melaleuca quinquenervia were, respectively, the most extensive and commonly recorded species in these families. ARIE Extent and decline of wetland types on the mid-north coast of New South Wales (modified from Goodrick, 1970)# Original 1969 Wetland type area (ha) area (ha) % Decline fresh meadows 18590 ( 2430) T9a0 seasonal fresh swamps ( 1340) semi-permanent fresh swamps 120 120 — open fresh waters* 40 40 - shallow saline lagoons 650 650 — teatree swamps 7490 6560 12.4 salt meadows 3520 3520 — reed swamps 2880 810 71.9) salt flats 160 160 mangrove swamps 1170 1170 = coastal bogs 1940 1940 - coastal Lepironia swamps 40 40 = 36600 18780 48.7 # estimates excluded two wetland types: shallow estuarine waters and sheoak swamps. * open fresh waters is the only floodplain wetland category identified by Goodrick as being dominantly open water, although relatively small areas of open water occur in other categories on the Macleay floodplain. Underlined categories are those occurring on the Macleay floodplain. TABLE 3 Numbers of native plant taxa in wetlands Taxa No. wetlands % Total no. > 30 4 0.9 26-30 3 0.7 21-25 9 Dall 16-20 31 12 11-15 101 23 Ea: 6-10 174 40.2 0-5 110 25.5 432 100.0 Twenty of the species recorded are introduced. They occupied about 180 ha or 1.5% of total wetland area at the time of the survey. The most extensive were Ezchhornia crassipes (94 ha), Salvinia molesta (33 ha) and Echinochloa crus-galli (31 ha). The first two mentioned are particularly troublesome floating weeds and had completely blanketed and infilled some wetlands. Cyperus brevifolius may be native to parts of northern Aus- tralia although it is considered to be introduced to the study area (K. Wilson, Natnl. Herb. N.S.W., pers. comm.). Four recorded species, Commelina cyanea, Cyperus polystachyos, Pseudognaphalium luteo- album and Ranunculus plebeius, are widely distributed in non-wetland habitats, particu- PROC. LINN. SOC. N.S.W., 111 (3), 1989 162 LOWER MACLEAY WETLANDS larly in higher rainfall areas. A further six species, Bacopa monniert, Bolboschoenus cald- welli, Juncus kraussu, Schoenoplectus litoralis, Sporobolus virginicus and Triglochin striata, are typical of brackish and estuarine habitats and occur only marginally in freshwater flood- plain wetlands. TABLE 4 Frequency of occurrence of plant taxa in wetlands according to Yo number of wetlands and % total wetland area (bracketed figures indicate number of introduced species in percentage classes) No. taxa No. taxa Percentage class (% no. wetlands) (% wetland area) 0-1.0 39(9) 107(20) 1.1-=2,,0 14(1) 1 2163-0 12(1) 3 371-420 4(2) = 4.1-5.0 4 1 5.1-6.0 (3) 1 6.1-7.0 5 = 7.1-8.0 3) = 8.1-9.0 4 2 9.1-10.0 2 2 10.1-20 12(2) 1 21-30 5(1) = 31-40 1 = 41-50 2(1) = 51-60 1 — 61-70 2 = 71-80 1 = 81-90 2 = 91-100 = — Four rare or uncommon plant species were recorded during the field survey: Cyperus platystylis (Cyperaceae): present in six wetlands over a total area of 0.1 ha; very rare in New South Wales although more common in northern Aus- tralia where it occurs on floating organic mats (K. Wilson, Natnl. Herb. N.S.W., pers. comm.); recorded only on mats of the introduced Salvinia molesta and Exchhornia crassipes; Eleocharis philippinensis (Cyperaceae): present in one wetland; third and southernmost record for the State; Maundia triglochinoides (Juncaginaceae): present in 30 wetlands and over a total area of 12.1 ha; known range only between Wyong and the Brisbane area (Aston, 1973; Sainty and Jacobs, 1981); considered rare on the New South Wales central coast by Beadle et al. (1982); generally rarely collected and of uncertain status (S. Jacobs, Natnl. Herb. N.S.W., pers. comm.); Potamogeton javanicus (Potamogetonaceae): present in one wetland; only known from the central and north coast regions of the State (Sainty and Jacobs, 1981) and recorded very rarely over this range although it 1s widespread in northern Australia and overseas (Aston, 1973); given as rare on the central coast by Beadle et al., (1982) and rarely collected from the Sydney region in recent years (R. Coveny, Natnl. Herb. N.S.W., pers. comm.). The majority of wetlands had dense, broad bands of herbaceous emergents around most of their perimeters (Table 5). The major fringing emergents, by far, were Persicarza hydropiper and Juncus polyanthemus x usitatus. Few wetlands had more than 10% of their margins lined with trees, the result of clearing for grazing. The commonest fringing trees were Casuarina glauca and Melaleuca quinquenervia. PROG. LINN. SOC. N.S.W., 111 (3), 1989 R. L. PRESSEY 163 TABLE 5 Summary of records for condition of marginal vegetation A B Percentage of perimeter with trees Percentage of perimeter with emergent vegetation Percentage No. wetlands % ‘Yotal no. Percentage No. wetlands % ‘Votal no. 91-100 1() Dos 91-100 at 75.0 66-90 12 2.8 66-90 19 4 at 36-65 15 32) 36-65 32 7.4 11-35 27 6.2 11-35 12 2.8 0-10 368 85.2 0-10 45 10.4 432 100.0 432 100.0 C. Average density and width of emergent vegetation Rating No. wetlands % Yotal No. 5 (very dense and wide) 316 T32 4 26 6.0 3 26 6.0) 2 19 4.4 1 (very sparse and narrow) 45 10.4 432 100.0 About 99% of the wetlands were grazed to some extent by cattle. At least five wet- lands had been separated from tidal influence by floodgates or earthen block banks. Drainage had directly affected 220 wetlands (51% of total number) and a wetland area of 12,236 ha (96% of total area). Additional, indirect effects of drainage on other wet- lands include separation into undrained sub-basins and could include local lowering of water tables. The actual hydrological effects of drainage would vary with factors such as the depth of drainage and of water tables and the size of catchments. In general, however, drained wetlands will have reduced storage capacity and will dry more readily com- pared to undrained ones. It 1s unlikely that any wetlands on the lower Macleay flood- plain have been completely eliminated by drainage, despite Goodrick’s (1970) estimates of wetland decline in the region (Table 2). His estimates are based on the elimination of value to waterfowl whereas wetlands were defined in this study by the occurrence of plants adapted to at least seasonal inundation. Many wetlands defined in this way are no longer of value as waterfowl habitat. Some 33% of the wetland area covered by the survey reported here was occupied by two taxa (Juncus polyanthemus x usitatus and Persicaria hydropiper) typical of marginal wetland conditions. Wetland drainage has probably been a major cause of the extensive distribution of these taxa. Two broad hydrological categories of wetlands were distinguished from the nature of their catchments. The first (Category 1) contains 342 wetlands (79% of total number) with a combined area of nearly 4800 ha (37% of total area) which have catchments extending no higher than the 10 m contour. They are filled only by groundwater and localized runoff from the surrounding alluvial flats and will generally have less stable water levels and will dry more readily than those in Category 2. The second type of wet- lands (Category 2), with additional drainages on the bedrock slopes surrounding the floodplain, number 90 (21% of total number) and have an overall area of about 8,000 ha (63% of total area). Most wetlands in Category 2 have catchments on bedrock smaller than 100 ha but the few large wetlands which make up most of the area of this type are fed by catchments PROG. LINN. SOG. N.S.W., 111 (3), 1989 164 LOWER MACLEAY WETLANDS larger than 1000 ha (Table 6). The catchment sizes for these wetlands are 1100 ha (East Kempsey Swamps, No. 275 in Fig. 1), 1790 ha (Belmore Swamp, No. 350), 5060 ha (Christmas Creek, No. 10) and 13,400 ha (Doughboy Swamp, No. 69). The other very large catchment covers 1880 ha and supplies wetland No. 249 which has an area of only about 1 ha. About 90% of wetlands in Category 2 have catchment area/wetland area ratios of 100 or less (Table 7). About 95% of the area in this type of wetland is fed by catchments that are at most 10 times larger than the individual wetlands. Larger catch- ments and larger ratios of catchment area/wetland area will generally confer greater permanence on wetlands or on particular parts of large, complex basins. Water levels in these wetlands would generally be more persistent because of the greater effectiveness of any rainfall in refilling the basins and perhaps because of greater base flow from ground- water in large catchments. TABLE 6 Catchment sizes for wetlands with catchments above the 10 m contour (Category 2) Catchment No. % Total Wetland % Total size (ha) wetlands no. Cat. 2 area (ha) area Cat. 2 0-10 43 47.8 67.9 0.9 11-100 29 32.2 78). 0.9 101-1000 13 14.4 503.0 6.3 > 1000 5 5.6 7340.8 Oil 90 100.0 7986.8 100.0 TABLE 7 Ratios of catchment area/wetland area for wetlands with catchments above the 10 m contour (Category 2) Catchment/ No. % Total Wetland % Total wetland ratio wetlands no. Cat. 2 area area Cat. 2 0-10 54 60.0 7553.8 94.6 11-100 28 31.1 ANT fl 52) 101-1000 7 7.8 14.5 0.2 > 1000 1 1.1 0.8 = 90 100.0 7986.8 100.0 Table 8 summarizes the occurrence of seven broad land uses in the catchments of the Macleay floodplain wetlands. Every wetland catchment contains some agricultural use and agriculture is the dominant or sole use in 93% of catchments. Areas of natural vegetation occur in 21% of catchments but occupy the largest parts of only 4% of catch- ments. None of the wetlands surveyed had completely natural catchments. Forestry, industrial areas, mining (only surface extraction of sand, gravel etc. in the case of the Macleay survey area), urban areas and waste disposal occurred in relatively few catch- ments and rarely or never occupied the largest proportions. The tenure of about 98% of the wetlands was solely or dominantly freehold. A small part of the Swan Pool (wetland No. 353) is within Hat Head National Park. The results reported here are estimates that are strictly valid only for the period of the field survey and for the dates of the aerial photography. The condition of wetlands and their catchments and the tenure of the wetlands will have since changed to some extent with new developments and changes in land use. In particular, short-term dynamics of wetland vegetation will have altered the occurrence of plant species and open water. Further surveys of the wetlands and research on the temporal variability of coastal freshwater wetlands are necessary to quantify and explain such changes. PROC. LINN. SOC. N.S.W., 111 (3), 1989 R. L. PRESSEY 165 TABLE 8 Land use occurrence in wetland catchments Present (% no. Dominant or sole Land use catchments) use (% no. catchments) Agriculture 100.0 93.3 Forestry 1.4 0.9 Industrial area 0.7 — Mining 2 Natural vegetation 2AES 4.2 Urban area 5.8 1.6 Waste disposal 1.4 ACKNOWLEDGEMENTS The study was funded by the N.S.W. National Parks and Wildlife Service as a consultancy. Phillippa Pressey provided much valuable assistance in the compilation and checking of the survey data and typed the original data base. Chris Ann Urquhart assisted in the preparation of the tables and drafted the figure. John Porter organized the analysis of the numerical distribution of open water categories and groups of plant taxa given in the Appendix. Staff of the National Herbarium of New South Wales are thanked for processing and identifying plant specimens. Thanks are due to Karen Wilson, in particular, for advice on identification. Karen Wilson, Surrey Jacobs and an anonymous referee provided helpful comments on the manuscript. References ASTON, H.I., 1973. — Aquatic Plants of Australia. Melbourne: Melbourne Univ. Press. BEADLE, N. C. W., Evans, O. D. and CAROLIN, R. C., 1982. — Flora of the Sydney Region. 3rd Edition. Sydney: Reed. BROOME, L. S., 1978. — Birds on north coast wetlands. Univ. New England Sch. Nat. Res. Rep. PR9. COASTAL COUNCIL OF NEW SOUTH WALES, 1985. — Coastal Wetlands of New South Wales. Department of Environment and Planning: Sydney. GooprRICck, G. N., 1970. — A survey of wetlands of coastal New South Wales. CSTRO Div. Wildl. Res. Techn. Memo. 5. Jacoss, S. W. L. and LaPINPpuRO, L., 1986. — Alterations to the census of New South Wales plants. Télopea 2: 705-714. , and PICKARD, J., 1981. — Plants of New South Wales: A Census of the Cycads, Conifers and Angiosperms. Government Printer: Sydney. MARGULES, C. R., NICHOLLS, A. O. and PREsSSEY, R. L., 1988. — Selecting networks of reserves to maximise biological diversity. Bzol. Cons., 43: 1-13. PRESSEY, R. L., 1981. — A Review of Literature on the Floodplain Wetlands of Coastal New South Wales. National Parks and Wildlife Service: Sydney. ——., 1987. — A Survey of Wetlands of the Lower Macleay Floodplain, New South Wales. National Parks and Wildlife Service: Sydney. ——, 1989. — Wetlands of the lower Clarence floodplain, northern coastal New South Wales. Proc. Linn. Soc. N.S.W,, 111: 143-155. , and MIDDLETON, M. J., 1982. — Impacts of flood mitigation works on coastal wetlands in New South Wales. Wetlands (Australia), 2: 27-44. SAINTy, G. R. and Jacoss, S. W. L., 1981 — Waterplants of New South Wales. Sydney: Water Resources Commission. WEsT, R. J., THOROGOOD, C. A., WALFORD, T. R. and WILLIAMS, R. J., 1985 — An estuarine inventory for New South Wales. N.S.W. Dept. Agric. Fisheries Bull. 2. PROC. LINN. SOG. N.S.W,, 111 (3), 1989 su Vb 88 su + 60 60 bE 6h 10 G9 GE 10 €°8 G°GZ 10 Z°8 8% su 10 aL el 9691 8°86 su 60 CW es Gal Gauci €°9G = 1-0 Val 69 < | ES 8°6 PESSI £16 = G0 8°19 0°69 Z su Gal 68 ec 8°8 QLITI GGL Q 3 G0 £89 Gia » su + (0) ss su Ih la 4 su + (om @) = su + Z0 su + (om) su + 20 su + L‘0 su + Z0 BOlIR potdn390 “Ou puepiom (ey)eore [Roy % [R101 % [R10], Co) WO ri (AO uoy) ssay saqoopua su “vy [() UDY) LazvaLe aguaLinz90 OU sayvIipur + oO 4 Goa 30 = x wo 23 50 ad ) yp x ow iS) (b) 1000 ~ uo Oo per licenced boat (kg) N uo uo (=) oO fo) Catch of Murray cod per licenced fisherman and c) : | 100 a 2 75 ° Ga 4 : 50 o ne po c oa o 5 oO Ta i 25 Ay 1940/41 1950/51 1960/61 1970/71 1980/81 Fig. 3. The total annual catch (a) and catch per licenced fisherman and per licenced boat (b) of Murray cod, and the percentage composition of Murray cod, golden perch and common carp (c) in the commercial catch from inland waters in N.S.W. between 1940/41 and 1983/84 (data from annual reports, N.S.W. State Fisheries). (b) O © catch per licenced fisherman (c) M——W@ = Murray cod A ------ A catch per licenced boat (J——U _-s golden perch => common carp PROG. LINN. SGC. N.S.W., 111 (3), 1989 S. J. ROWLAND 207 1940/41 and 1955/56, but then the total catch and the catch per licenced fisherman declined dramatically (Fig. 3a, b). Murray cod was the major species in the inland fishery until 1951/52, and between 1940 and 1951 cod comprised 42-65% of the total annual catch from inland N.S.W. However, after 1951/52, golden perch (Macquaria ambigua) replaced Murray cod as the major native species in the N.S.W. inland fishery (Fig. 3c). The total catch, catch per licenced fisherman and the percentage composition data all strongly suggest that there was a dramatic decline in the abundance of M. peeli between 1955 and 1964. There was a concurrent decline in the commercial catch of Murray cod from the Murray River in South Australia (Reynolds, 1976). Between 1940/41 and 1962/63 the annual total catch of Murray cod in N.S.W. exceeded 35,000 kg; however since 1963/64, with the exception of 1974/75, the annual catch has remained below 30,000 kg and the catch per licenced fisherman below 200 kg. The continued small catch and low catch per licenced fisherman of Murray cod in the N.S.W. commercial fishery since 1960/61 (Fig. 3) indicate that the fishery remains in a depressed state. These catch statistics also indicate that there has been no major reduction or increase in the abundance of Murray cod since the mid 1960's. Consequently, it appears that stocks of Murray cod have remained stable, but at relatively low levels over the last 25 years. Reduced Distribution There has also been a reduction in the distribution of M. peels. Many historical reports indicate that the species was common in rivers where cod are now rare or no longer found. The explorer George Evans, the first European man to see the species (Stanbury and Phipps, 1980), observed cod in the Fish River before he reached the present site of Bathurst. The type locality of M. peeli is the Peel River, N.S.W. probably near the present site of Tamworth (Mitchell, 1838) and in 1836 an enormous 120 lb ‘River codfish of the Colonists’ was found entangled and struggling near the bank in a pond of the Cudegong River (Bennett, 1864). Murray cod are now extremely rare in the Fish, Peel and Cudegong rivers. In Victoria, Murray cod abounded in the Loddon, Campaspe and Goulburn rivers and their tributaries, even where the waters ‘dwindled into the most insignificant streams’ (Wilson, 1857). By the late 1940’s the populations of Murray cod and other native fishes had declined in these rivers (Langtry, in Cadwallader, 1977) and there are now very few localities in Victoria where Murray cod can be considered common (Cadwallader and Backhouse, 1983). Fisheries Management and Research Concern about the stocks of Murray cod was expressed as early as the 1880 Royal Commission enquiring into the Fisheries of N.S.W. (Macleay et al., 1880) and from 1883 until about 1895 there was some supervision of inland waters (Dakin and Kesteven, 1938). Dannevig (1903) detailed measures for restricting gear, protecting fry and young fish and the imposition of a closed season. From 1905 to 1910 serious attempts were made to improve the cod fishery, including experiments conducted in 1905 by H. C. Dannevig on the artificial propagation of cod (Farnell, 1906; Dakin and Kesteven, 1938). Whitley (1937) briefly discussed the available information on the distribution, fishery, breeding and taxonomy of Murray cod and included a complete bibliography containing mostly taxonomic references. In 1936, a conference on the Murray River fisheries, attended by representatives from N.S.W., Victoria and South Australia, adopted a closed season of September, October and November for the taking of Murray cod; set minimum legal lengths; suggested that hatcheries be established; asked the Murray River Commission to construct a fishway at Lock 15 at Euston; and suggested PROG. LINN. SOG. N.S.W,, 111 (3), 1989 208 MURRAY COD HISTORY AND FISHERY that the breeding habits and migration of freshwater indigenous fishes be studied by each State (Isherwood, 1939). However, since that time only five studies have contributed significantly to the knowledge of the biology of M. peeli. Dakin and Kesteven (1938) presented brief notes on the natural history, behaviour in captivity and the spawning season of Murray cod; they also discussed the cod fishery including the evidence for, and possible causes of the decline of cod stocks. Dakin and Kesteven artifically bred cod by capturing and strip- ping ripe fish; the eggs, embryonic development, larvae and fry were described. However, this part of the study was restricted by the difficulty of procuring ripe brood- fish from the wild. The need for future research into the biology and breeding of this species was emphasized. J. O. Langtry conducted an ecological survey of the Murray River and some of its tributaries in 1949-50. Unfortunately, his report was not published, and it wasn’t until 1977 that the manuscript was rewritten and presented by Cadwallader (1977). The report contains data on the relative abundance of fishes in the study area and describes the differences between Murray cod and trout cod (Maccullochella macquariensis). Langtry also made observations and collected some quantitative data on the distribution, diet, breeding biology and growth rate of Murray cod. During the 1960’s, John Lake studied the reproductive biology of native fishes at the Inland Fisheries Research Station, Narrandera, N.S.W. and his research demon- strated that critical temperatures and rising water levels in ponds (and presumably a flood or fresh in the wild) triggered the spawning of some species (Lake, 1967 a,b). Cadwallader and Gooley (1985) collected data on the spawning of Murray cod in earthen ponds, and developed techniques for the artificial propagation and rearing of M. peeli. Rowland (1985) conducted research into the biology and artificial breeding of M. peeli, and some of his findings form the basis of the following discussion. POSSIBLE CAUSES OF THE DECLINE Reduced Larval Recruitment The high survival of fish larvae is dependent on the availability of relatively high concentrations of suitable-sized food organisms at the commencement of exogenous feeding; suboptimal feeding conditions generally result in death due to starvation or predation (May, 1974; Pitcher and Hart, 1982). Hjort (1926) hypothesized that the degree of mortality of larvae during a ‘critical period’ after the completion of yolk sac absorption determined the strength of year-classes in natural populations. Although the relationship between larval mortality during the ‘critical period’ and year-class strength is difficult to determine in nature (May, 1974) it is generally thought that the survival rate of fish larvae is the most important factor determining the strength of year-classes (Beverton, 1962; Gulland, 1965; May, 1974; Pitcher and Hart, 1982). Many overseas studies have shown that strong year-classes of freshwater fishes are established when the breeding season coincides with rising or high water levels (e.g. Agegus and Elliott, 1975; Stevens, 1977; Marshall, 1982; Beam, 1983). Although M. peel: spawned annually in the southern tributaries of the Murray-Darling river system between 1977 and 1980, relatively strong year-classes were only established when the breeding seasons coincided with high river levels or floods (Rowland, 1985), demon- strating that floods in October and November provide optimum conditions for the survival and recruitment of M. peeli larvae. The floodplain areas of the Murray-Darling river system are highly productive. When they are inundated in spring or summer, a rich source of terrestrial nutrients, plus the plankton, aquatic insects (in particular chironomid larvae) and other organisms PROC. LINN. SOG. N.S.W., 111 (3), 1989 S. J. ROWLAND 209 of the billabongs become available to the aquatic community of the rivers (Frith, 1959; Shiel, 1980; Maher and Carpenter, 1984). Zooplankton, chironomid larvae and other aquatic insects are the major food items of the larvae and fry of Murray cod and golden perch in earthen ponds, and a delay of several days in the availability of food to larvae after the completion of yolk sac absorption results in reduced survival in both species (Rowland, 1985, 1986). The construction of dams, high-level weirs and levee banks on the major tributaries of the Murray-Darling system has altered the natural flow and temperature regimes and dramatically reduced the frequency, extent and duration of floods (Lake, i971; Reynolds, 1976; Cadwallader, 1978; Walker et al., 1978; Walker, 1979). The Murray River no longer floods annually in spring and much of the vast anabranch, billabong and floodplain areas of the Murray and Murrumbidgee rivers have been eliminated and do not flood except under extraordinary circumstances (Langtry, in Cadwallader, 1977; Shiel, 1980). Consequently optimum conditions for the survival of the larvae of Murray cod, which usually spawn in October or November, now rarely occur. It is suggested that the reduced frequency, extent and duration of spring flooding in the Murray-Darling river system has led to an overall reduction in larval recruitment and that this is a major cause of the decline in the abundance of M. peelz. This hypothesis is supported by the change in the relative proportion (by weight) of Murray cod and golden perch in the commercial catch from inland N.S.W. (Fig. 3c). Prior to 1951/52, M. peel: was the major species; however, since then golden perch have formed a much greater percentage of the annual catch than have Murray cod. Golden perch require a substantial rise in water level, when temperatures are about 23°C, to induce spawning (Lake, 1967a); if suitable conditions do not occur in the wild, adults remain at an advanced stage of gonadal development until March or April (Mackay, 1973). Therefore M. ambigua can delay spawning over a six month period whereas M. peelt spawns only during spring and early summer when the water temperature is about 20°C (Rowland, 1983, 1985; Cadwallader and Gooley, 1985). The reduced frequency and extent of flooding, in particular the reduction of flooding which usually occurred each spring in the Murray and Murrumbidgee rivers (Langtry, in Cadwallader, 1977; Walker e¢ al., 1978; Walker, 1983) would be expected to affect larval recruitment to a greater extent in M. peel: than in M. ambigua. The major impoundments on the Murray-Darling river system in N.S.W. were constructed between 1907 and 1976; Burrinjuck Dam 1907 (enlarged 1957), Hume Dam 1936 (enlarged 1961), Wyangla Dam 1936 (enlarged 1971), Yarrawonga Weir 1939, Keepit Dam 1960, Menindee Lake Storage Scheme 1960, Burrendong Dam 1967, Blowering Dam 1968, Pindari Dam 1969, Copeton Dam 1976 (Anon., undated; Walker, 1980). By 1950, the effects of Hume Dam and Yarrawonga Weir on the flow in the Murray River were clearly evident (Langtry, in Cadwallader, 1977). The cumulative effects of the major impoundments and water storage schemes would have been substantial by 1960 when there was an apparent rapid decline in the abundance of Murray cod. Overfishing The relatively large inland commercial fishery which existed between the mid 1800’s and the late 1930’s and which was based primarily on Murray cod (Macleay et al., 1880; Dannevig, 1903; Stead, 1903; Dakin and Kesteven, 1938) would have placed intense fishing pressure on cod populations. Previously unfished populations of long- lived fishes consisting of 12 or more year-classes are extremely susceptible to exploi- tation; and with unchanging recruitment the absolute size of the total stock will decline markedly, or even catastrophically under moderate exploitation (Ricker, 1963). It is PROG. LINN. SOG. N.S.W., 111 (3), 1989 210 MURRAY COD HISTORY AND FISHERY therefore probable that the decline in abundance of Murray cod, at least until the 1930's, by which time the inland commercial fishery had become unprofitable for large-scale operators (Pollard and Scott, 1966), was caused primarily by overfishing. A possible reduction in the exploitation of cod populations after the 1930’s due to the depressed state of the fishery, the Depression and World War II, may have resulted in numbers and the catch per licenced fisherman/boat remaining reasonably stable or even increasing (Fig. 3) until the late 1950’s when, as suggested, the effects of reduced larval recruitment became apparent by the rapidly declining stocks. It is also possible that the recreational fishery which developed in the 1950’s (Anon., 1956; Poole, 1984) contributed to the decline of cod stocks during this period. English Perch (Redfin) English perch, Perca fluviatilis, were abundant and sympatric with M. peel: in the southern tributaries of the Murray-Darling system between the late 1940’s and the 1960’s. Langtry (in Cadwallader, 1977) found that the diet of P fluviatilis was identical to that of M. peeli and M. ambigua, and small fishes become a major part of the diet of larger English perch (Lake, 1967c). English perch larvae and juveniles feed on zooplankton, crustaceans and insect larvae, and because P. fluviatilis spawns in early spring when temperatures are about 12°C; usually the last week or so in August in southern N.S.W. (Lake, 1967c), juvenile English perch may prey on, and possibly compete for food with Murray cod larvae and fry, particularly during drought periods when food resources are limited. The catch data on fishes from the Kerang Lakes, Victoria, between 1919 and 1949 (Figure 1 in Cadwallader, 1977) demonstrated that when English perch were abundant, native fish were scarce and vice versa. It is therefore possible that English perch have contributed to the decline of Murray cod in the southern parts of the Murray-Darling river system. Other Factors Factors such as siltation, desnagging, channelization, depressed water tempera- tures below impoundments, and barriers preventing spawning migrations have been implicated in the decline of native fishes (see Merrick and Schmida, 1984). While some of these factors may be responsible for the decline of some species in certain areas, because of the great size of the Murray-Darling river system it is unlikely that they have been major contributing factors to the decline of M. peels which is widely distributed throughout the system (Lake, 1971) and does not undergo extensive migrations (Reynolds, 1983). SUMMARY Historical aspects of Australia’s most famous and largest inland freshwater fish, the Murray cod, Maccullochella peeli, are briefly reviewed. Fossil records suggest that ancestral Murray cod are of, at least Miocene age. The Murray cod played a prominent part in the mythology and culture of some aboriginal tribes. Explorers, including George Evans, John Oxley, Thomas Mitchell, and Charles Sturt, and the early, inland settlers were astounded by the abundance, size and delicacy of Murray cod. The species was held in such high esteem that it was considered worthy of acclimatization in England and Europe, and although this did not eventuate M. peel: was extensively trans- located within and outside its natural range of the Murray-Darling river system during both the 19th and 20th centuries. During the late 1800's a large, inland, commercial fishery developed. It was based on Murray cod and was located mainly on the mid and lower reaches of the Murray and PROC. LINN. SOC. N.S.W., 111 (3), 1989 S. J. ROWLAND 211 Murrumbidgee rivers. Historical records and catch statistics indicate that there has been a dramatic decline in the abundance and a reduction in the distribution of M. peelz. Possible causes of the decline are briefly discussed. It is suggested that overfishing contributed to a decline between the late 1800’s and the 1930's, but that the reduced frequency, extent and duration of spring flooding in the Murray-Darling river system, caused by the construction of dams, high-level weirs and levee banks, has adversely affected larval recruitment in M. peeli. This has caused the dramatic decline in abundance of cod during the 1950’s and the maintenance of stocks at relatively low levels over the last 25 years. ACKNOWLEDGEMENTS I gratefully acknowledge Ray Gregg for providing the photograph of the paddle- steamer ‘Mayflower’. I express my appreciation to Peter Williamson and David Rodgers for preparing the figure and photograph respectively, and to Margaret Leedow and Denis Reid for providing the fisheries catch statistics. I thank Barbara Butler for typing the manuscript. References AGGuS, L. R., and ELLIOTT, G. V., 1975. — Effects of cover and food on year-class strength of largemouth bass. 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OXLEY, J., 1820. — Journals of Two Expeditions into the Interior of New South Wales, undertaken by Order of the British Government in the Years 1817-18. London: John Murray. PHILLIPS, N. C., 1863. — The Murray River cod. Sydney Morning Herald, 24 Sept. PITCHER, T. J., and Harr, P. J. B., 1982. — Fisherves Ecology. Connecticut: A.V.I. POLLARD, D. A., and ScorT, T. D., 1966. — River and reef. In, MARSHALL, A. J., (ed.) The Great Extermin- ation. Melbourne: William Heinemann. POOLE, D. E., 1984. — Inland fishery on River Murray — some historical facts. SAFIC 8: 20-22. RAMSAY SMITH, W., 1930. — Myths and Legends of the Australian Aboriginals. London: Harrap. REYNOLDS, L. F., 1976. — Decline of native fish species in the River Murray. SAFIC 8: 19-24. ——,, 1983. — Migration patterns of five fish species in the Murray-Darling river system. Aust. J. Mar. Freshw. Res. 34: 857-871. RICKER, W. E., 1963. — Big effects from small causes: two examples from fish population dynamics. j. Fish. Res. Board Can. 20: 257-264. ROWLAND, S. J., 1983. — Spawning of the Australian freshwater fish Murray cod, Maccullochella peeli (Mitchell), in earthen ponds. J. Fish. Biol. 23: 525-534. PROC. LINN. SOC. N.S.W., 111 (3), 1989 S. J. ROWLAND Dales) —., 1985. — Aspects of the biology and artificial breeding of the Murray cod, Maccullochella peeli and the eastern freshwater cod, M. tkern. sp. North Ryde, N.S.W.: Macquarie University, Ph.D. thesis, unpubl. , 1986 — The hormone-induced spawning and larval rearing of Australian native freshwater fish, with particular emphasis on the golden perch, Macquaria ambigua. In, REYNOLDS, L. F., (ed.), Proceedings of the First Australian Freshwater Aquaculture Workshop. N.S.W. Sydney: Department of Agriculture. SHIEL, R. J., 1980. — Billabongs of the Murray-Darling system. /n, WILLIAMS, W. D., (ed.), An Ecological Basis for Water Resource Management. Canberra: Australian National University Press. STANBURY, P., and PHIpPs, G., 1980. — Australia’s Animals Discovered. Sydney: Pergamon Press. STEAD, D. G., 1903. — A description of gear used on the Murray; with notes on the natural enemies of the Murray cod, and on the various fishes, crustacea, etc. captured or mentioned. Jn, Murray Cod Fisheries. Sydney: Government Printer. STEVENS, D. E., 1977. — Striped bass (Morone saxatilis) year class strength in relation to river flow in the Sacramento-San Joaquin estuary, California. Trans. Am. Fish. Soc. 106: 34-42. Sturt, N. G., 1899. — Life of Charles Sturt. Sometimes Capt. 39th Regt. and Australian Explorer. London: Smith, Elder and Co. TayLor, G., WALKER, D. H., JONES, N.O. and Hutka, J., 1980. — The nature and significance of lacustrine deposits near Bunyon, N.S.W. Abstract of Symposium on Cainozoic Evolution of Continental South west Australia, Canberra, November 1980. TINDALE, N. B., 1951. — Aboriginal net making. 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Qld Mus. 11: 113-148. ——, 1955. — The largest (and the smallest) Australasian fishes. Aust. Mus. Mag. 11: 329-332. , 1959. — The freshwater fishes of Australia. In, KEAST, A., CROCKER, R. L. and CHRISTIAN, C. S., (eds.), Biogeography and Ecology in Australia. The Hague: W. Junk. WILSON, E., 1857. — On the Murray River Cod, with particulars of experiments instituted for introducing this fish into the River Yarra-Yarra. Trans. Phil. Inst. Vic. 2: 23-34. PROC. LINN. SOC. N.S.W,, 111 (3), 1989 IS na! halt Bly ' . aia ye ais f hele may Lg Microhydromys musseri n. sp., a New Murid (Mammalia) from the ‘Torricelli Mountains, Papua New Guinea T. F. FLANNERY FLANNERY, T. F. Microhydromys musseri n. sp., a new murid (Mammalia) from the Torricelli Mountains, Papua New Guinea. Proc. Linn. Soc. N.S.W. 111 (3), 1989: 215-222. Microhydromys mussert n. sp. differs greatly from the only previously described species of Microhydromys (M. richardsonz) in its larger size, brown dorsum with a markedly contrasting venter, broader skull, palate morphology, and dentition details. Both species however share synapomorphies not seen in other hydromyine murids. Micro- hydromys mussert n. sp. is thus far known from a single specimen collected near the summit of Mt Somoro, Torricelli Mountains (part of the North Coast Ranges). It is the third mammal species endemic to these ranges to be described. Three additional speci- mens of Microhydromys richardsoni, which was previously known only from the holotype, are described. These add significantly to knowledge of the species distribution and morphology. T. F. Flannery, Australian Museum, Sydney, Australia 2000; manuscript received 31 January 1989, accepted for publication 19 April 1989. KEY WORDS: Microhydromys mussert, New Guinea, Hydromyinae, zoogeography. INTRODUCTION The murid subfamily Hydromyinae consists of 10 genera in the Australo-Papuan region. Thomas (1898) and Misonne (1969) have referred Asian taxa to the Hydro- myinae, but Musser (1982) notes that it is not yet clear if any of these taxa do indeed represent part of the hydromyine radiation. All of the Asian taxa are readily distinguish- able from members of the Australo-Papuan group and none resemble the species of Mucrohydromys. For this reason, and because of their uncertain status, I have restricted comparisons of the new species to Australo- Papuan taxa. Within the New Guinean Hydromyinae are a group of six species, placed in four genera, that are known as microhydromyines. They are small, shrew-like murids with short, sleek coats, tiny eyes and pointed snouts. Included are some of the rarest and least known Melanesian mammals. The genera Mayermys and Neohydromys are monotypic, while Pseudohydromys includes two similar, probably parapatric species. Microhydromys includes two species that differ greatly in morphology, one of which is described here as new. Previous taxonomic work upon microhydromyines has been limited, and it is not yet clear whether these species represent a monophyletic group, or are simply pheneti- cally similar and paraphyletic. Likewise, the generic limits are in some cases poorly understood, and it is possible that in future some genera will be synonymized. Microhydromys richardsoni Yate and Archbold, 1941 was described on the basis of a single adult male taken at an altitude of 850m near Bernhard Camp, Idenburg River, Irian Jaya. It is one of the more distinctive microhydromyines, possessing grooved upper incisors and a broad skull with a short rostrum. A brief mention of the existence of an additional specimen (Menzies and Dennis, 1979) constitutes the only other reported occurrence of this species. In this work a new species of Microhydromys, from the North Coast Ranges, is described; descriptions of 3 additional specimens of M. richardsoni are provided. Although the new species is known only from a single specimen its description as a new PROC. LINN. SOC. N.S.W., 111 (3), 1989 216 A NEW MURID FROM PAPUA NEW GUINEA taxon is considered justified on the basis of its highly distinctive morphology. The likeli- hood of additional material being found in the near future is also low. MATERIALS AND METHODS Dental terminology follows Musser (1981), and colours where capitalized follow Smithe (1974). Measurements are in millimitres and weights in grams. Abbreviations are as follows: AM M, Australian Museum mammal specimen; AMNH, American Museum of Natural History mammal specimen; BBM, Bishop Museum mammal specimen. SYSTEMATICS Maucrohydromys ‘Yate and Archbold, 1941 Maicrohydromys mussert n. sp (Figs 1-2, Table 1) Holotype and Type Locality: BBM101737, puppet skin and skull with dentaries of an adult male, collected on 12 November 1972 by A. B. Mirza at 1,350m on Mt Somoro (3° 22'S, 142° 09’E), Torricelli Mountains, West Sepik Province, Papua New Guinea. Etymology: For Dr Guy Musser, who has contributed so abundantly to systematic research on the murids of Asia and Melanesia, and who has so greatly increased our understanding of the shrew-like murids of the region. Diagnosis: Microhydromys musseri n. sp. can be distinguished from M. richardson: in the following ways: it is larger (Table 1); the dorsum is Ochre Brown and the sharply demar- cated venter Cinnamon (as opposed to the nearly uniform grey colouration of M. richard- soni); there is no longitudinal groove on the upper incisors; the palate is more deeply concave; the skull is absolutely and relatively broader; I/1 is less procumbent; a posterior cingulum is present on M/1. Description: Fur short and dense, tail and ears appearing naked to the unaided eye. Dorsum near Ochre Brown, grading to Cinnamon Brown on flanks. Venter sharply demarcated and Cinnamon with two small, irregular white patches on thorax. Throat Cinnamon, crown of head Fuscous. Eyes surrounded by a circle of blackish hairs. Most vibrissae pale, but a few black; similar in length, distribution and density to those of other microhydromyines such as M. richardsont and N. fuscus. Hands and feet thinly furred with pale hairs on dorsal side. Tail slightly paler below than above and mottled with light patches becoming prominent distally. Flesh of ears dark. Tail scales ill-defined but form regular rings with a single hair per tail scale. Flesh of the hands and feet shrivelled, but enough can be seen to determine that the pads of the forefeet were striated. Tiny rhinarium naked, toes unwebbed. Upper incisors ungrooved and enamel orange. Lower incisors with paler orange enamel and less procumbent than in most other hydromyines. Molars heavily worn, with much crown detail lost. M1/ elongate, narrow, subrectangular. T2 large, obvious, much worn. A slight groove on the anterolingual face separates Tl and T2. Distinct enamel basin separates anterior and median lophs. Wear has obliterated almost all detail distal to this point. However, a slight groove discernable between median and posterior lophs on buccal face. T4 present as well-developed ridge. All M2/ crown details obliterated by wear. M2/ triangular in outline, with apex facing posteriorly. M/1 heavily worn. Protolophid horseshoe-shaped dentine basin surrounded by remnant of enamel crown. Hypolophid similar, but subrectangular shaped basin. Posterior cingu- PROC. LINN. SOC. N.S.W., 111 (3), 1989 T. F FLANNERY Daly) Fig. I. The crania of A, Microhydromys musseri (holotype) BBM 101737. B, BBM 60202, M. richardsoni. C, AM M14166, M. richardsonz. lum large and ovoid, positioned at anterolingual margin of tooth. M/2 smaller than M/1. Protolophid subrectangular and heavily worn basin. Hypolophid crescent shaped basin. Lophids separated by simple interlophid valley. Cranium slightly damaged; left occipital condyle sheared off and basoccipital shattered then reglued. Rostrum shorter than in other microhydromyines, and broadened posteriorly. Premaxillae do not extend beyond incisors. Masseteric foramen lunette shaped and sloping anterodorsally, but with vertical orientation at posterior end. Infraorbital foramen moderate in size, not expanded as in Hydromys, Parahydromys and Crossomys species. Incisive foramina short and bowed, posterior ends located just posteromedial to zygomatic plates. Zygomatic plates broad and flared laterally. Zygomatic arch thin. Braincase moderately inflated. Bregmatic anteroposteriorly very short but broad. Palate deeply excavated, with molar alveolar margin raised into a ridge which extends well ventral to palate floor. Palatal foramina partially obscured by over- hang of this ridge. Well-defined grooves run from palatal to incisive foramina. Left pterygoid broken away. Otic notch large, bulla unexpanded. PROG. LINN. SOG. N.S.W., 111 (3), 1989 218 A NEW MURID FROM PAPUA NEW GUINEA Fig. 2. Skins of A, M. richardson: (AM M14166) B, ventral view and C, dorsal view of M. richardsoni (BBM60202), D, ventral view and E, dorsal view of Microhydromys musseri n. sp. (holotype) BBM101737. DISCUSSION Systematics The assignation of the newly described species to the genus Microhydromys is based upon a cladistic interpretation of aspects of hydromyine morphology. Phenetically, mussert n. sp. and M. richardsoni appear to be rather different. However they share a number of key, apparently derived, features that are either rare or are not seen elsewhere among the Hydromyinae. These include the following. 1) The M1/is extremely narrow and elongate, and more nearly rectangular in occlusal outline than in other hydromyines. This feature is unique in the subfamily and is not seen among other Australo-Papuan murids. Thus on the basis of outgroup comparison this feature 1s likely to be derived. 2) The lingual end of the interloph valley between the anterior and medial lophs of M1/ is very narrow, and partially closed off by a low ridge from T1. The entire lingual PROC. LINN. SOC. N.S.W., 111 (3), 1989 T. F FLANNERY 219 margin of the tooth thus forms a continuous linear face which is parallel with the skull midline. Again, this feature is not seen in other hydromyines and is doubtless derived. 3) The rostrum is short relative to skull length and yet the interorbital canals are not greatly enlarged. The only other hydromyines with such a short rostrum are the species of Hydromys, Crossomys and Parahydromys (here called the ‘Hydromys group’). These very large hydromyines however all possess greatly enlarged infraorbital canals. Most murids (including the plesiomorphic hydromyine Leptomys elegans) have longer rostra. Thus it seems likely that the presence of a short skull is a derived feature. Because in other aspects the morphology of members of the ‘Hydromys group and the species of Micro- hydromys is so different it seems likely that this similarity is due to convergence. 4) The palatal foramina are partially closed posteriorly and are narrow. Again, these features are unique in the subfamily and on the basis of outgroup comparison are likely to be derived. Other aspects of morphology of the two Maicrohydromys species are however not shared between these taxa, but are either unique, or are shared by one or the other species with other hydromyines. These features are interpreted as being either retained symplesiomorphies or as being due to convergence; the features are as follows. The pelage of M. mussert n. sp. is bright. Its warm brown and cinnamon tones and sharply contrasting venter are not approached in any other microhydromyines. Among other hydromyines, only the species of Leptomys and one species of Paraleptomys possess such colouration. The species of Leptomys are particularly plesiomorphic hydromyines in many aspects of their morphology, and no other features would suggest a relationship with Microhydromys. Thus this feature may well be due to either convergence or else be a retained symplesiomorphy. Another striking feature of /. musseri n. sp. that is most closely approached only in apparently distantly related taxa is the extremely concave palate. Only in Xeromys myoides, among other hydromyines, is the condition seen in M. musseri n. sp. approached. However, here details of actual palatal structure differ, suggesting that palatal concavity in these taxa is the result of convergence. The palatal foramina in X. myozdes are not over- hung by the ridges supporting the cheekteeth as in M. mussern n. sp., but are subovate and open. Furthermore, in cross section the palate of M. mussert is more angular than in X. myoides (where the cross section is arch shaped). Overall skull shape at first suggested that a close relationship might exist between M. musserin. sp. and the ‘Hydromys group’ species, for these taxa have a short rostrum and are superficially similar in skull shape. However, in species of the ‘Hydromys group’ T1 and T4 on M1i/ are not extended posteriorly as they are in other New Guinean hydromyines. Posterior extension of these cusps must be regarded as derived, as it is not seen in plesiomorphic murid groups. This suggests that all hydromyines except members of the ‘Hydromys group may be monophyletic, and that similarities between M. musseri n. sp. and the species of Hydromys are either plesiomorphic or due to convergence. A final striking feature of M. musseri n. sp. is the presence of a distinct and large posterior cingulum on M/1. Only in the species of Leptomys, Paraleptomys and Neohydromys among hydromyines is this structure otherwise retained. The widespread presence of a posterior cingulum on the lower molars of other murid groups (including all non- hydromyine Australo-Papuan taxa) suggests that the loss of this structure is derived. That the posterior cingulum appears to have been retained in several unrelated hydromyine taxa indicates that its loss within the subfamily has occurred independently in several lineages. Mucrohydromys richardsoni possesses one feature that is unique in the Hydromyinae. This is the presence of longitudinally grooved incisors. Such grooved incisors are, however, seen in a number of other murid taxa, such as the species of Mylomys, Pelomys PROC. LINN. SOC. N.S.W., 111 (3), 1989 220 A NEW MURID FROM PAPUA NEW GUINEA and Golunda. In Vandeleurta oleracea the incisors are only occasionally grooved, while in Mus xenodontus only one incisor is grooved. Misonne (1969) suggests that this indicates that the presence of grooves on the upper incisors of murids is under relatively simple genetic control and that it has only limited systematic importance. These data, together with the fact that the grooved incisors of M. mchardsoni are unique (autapomorphic) among near relatives, suggests that not too much importance should be placed upon this feature in phylogenetic analysis. In summary, it would appear that M. musseri n. sp. and M. rchardsoni are each other’s nearest relatives, but that both possess a number of independent specializations not seen in the other. This may in turn indicate that these taxa have been evolving separately for a considerable period of time. On this basis, a case could be made for either placing each in a monotypic genus, or retaining both in Microhydromys. The latter course has been chosen because I feel that the recognition of yet another monotypic genus of microhydromyine murid is not desirable until relationships within the group are clarified; furthermore such a decision would not indicate the nature of the relationship between M. richardsoni and M. musseri n. sp. Biology and Zoogeography A note on the label of the type specimen of M. musseri n. sp. indicates that it was taken at an altitude of 1,350m in a snap trap in ‘forest’. At altitudes above about 1,200m Mt Somoro and the other high peaks of the North Coast Ranges are covered by mossy forest which differs sharply from the forest at lower altitudes. It is now becoming clear that these forests harbour a distinctive mammal fauna unlike that found anywhere else in New Guinea. The first endemic mammal to be described from these ranges (in this case the Cyclops Mountains) was also a hydromyine, Faraleptomys rufilatus Osgood, 1945. It has subsequently been found in the Mt Somoro area. Zeigler (1981) described a second endemic, the large petaurid Petaurus abidi from Mt Somoro. This paper describes a third endemic, which is yet another hydromyine. Furthermore, recent fieldwork undertaken in the Somoro area has revealed the presence of additional, as yet undes- cribed, endemic mammal species. In addition to these taxa, a number of slightly more widespread species are found in this habitat. These include Pseudocheirops albertisi1, also known from the Vogelkop and Japen Island, and Dendrolagus inustus, which has a similar distribution. The area of mossy forest available to these species in the Torricelli Moun- tains is tiny (ca. 39 km?), but in the recent past was probably larger, when Ice Age cooling depressed altitudinal zones. However, it appears that at no time in the last few million years was this region in contact with the mossy forests of the Central Cordillera. This is reflected in the ‘unbalanced’ assemblage of species in this habitat. For example, no endemic dasyurids are known from these ranges, while the Central Cordillera has 6 species found exclusively at mid-high altitudes. Perhaps the fact that two out of the three described endemic mammals from the North Coast Ranges are insectivorous hydromyines reflects this lack. In the absence of the insectivores and carnivores which inhabit this forest type elsewhere, it may well have been that the hydromyines that could invade the habitat underwent a modest radiation. Whatever the case, it is apparent that no similar event happened elsewhere, as no endemic hydromyine species have been found on other isolated mountain ranges in New Guinea. Additional Specimens The only detailed information previously published regarding Muocrohydromys richardsoni is that contained in the type description and paraphrases of it. The holotype is an adult male, taken, presumably in a snap trap (the posterior part of the skull is broken in a way that suggests snap trap damage), at an altitude of 850m 4km SW of Bernhard PROC. LINN. SOC. N.S.W,, 111 (3), 1989 T. F. FLANNERY 221 camp on the Idenburg River, Irian Jaya on 16 March 1939. Only three further specimens have been collected since that time, and two of these have been examined for this study. The first collected is an adult male (AMNH 198790). This was snap trapped under a small log in secondary growth at an altitude of 670m at Wanuma in the Adelbert Mountains, Madang Province by A. C. Zeigler on 21 October 1967. I have been unable to examine this specimen but Dr G. Musser kindly made the following notes for me; the terminal 35mm of the tail is mottled white, and the skull is so shattered that measure- ments could not be made. The Adelbert Range is a small, relatively low mountain range (most of it barely exceeding 1,000m) which is isolated from the New Guinean Central Cordillera. The second additional specimen to be collected 1s another adult male (BBM60202). This was snap trapped in an area of secondary forest-Eucalypt savannah near Sirinumu Dam, Sogeri area, Central Province Papua New Guinea on 28 October 1968. Sirinumu Dam lies at an altitude of ¢. 550m. This is a most unexpected locality, as the Idenburg River and Adelbert Range specimens must have come from much wetter forest. The measurements of this specimen closely approach those of the holotype for the most part (Table 1). The pelage and cranial morphology are also similar to that of the holotype. They differ primarily in that: approximately 37mm of the distal part of the tail is white tipped as opposed to 10mm in the holotype; the grey of the dorsum is interrupted by irregular and inconspicuous ginger blotching (the individual blotches being 1-2mm in diameter). TABLE 1 Measurements for the 5 known specimens of Microhydromys. Measurements for AMNH 152079 are from Tate (1951). Measurements in mm and weight in grams. 1 = length, w = width, inc. = incisive M. mussert M. richardsoni Measurement BBM-NG AMNH AMNH AMM BBM-NG /Weight 101737 198790 152079 14166 60202 head body 1. 108 83 80 86 86 tail vent 1. 101 84 92 79 83 hindfoot (su) 1. 22 19 20 = 19 ear (n) 1. 13 11 8 = 12 weight — 9 = 12 = condylobasal 1. 23.6 = = 20.2 19.6 bizygomatic w. i133}.40 = Q2 10.1 9.6 palate 1. 13.6 = 10.0 9.5 Q7 rostral w. 5.7 — = 4.1 4.0 inc. foramen 1. DP = ES) 2.1 1.9 interorbital w. D2 — 4.4 4.6 4.2 nasal 1. 8.0 = 6.4 7.0 5.9 maximum nasal W. 3.0 - 2.6 2.6 223 mastoid w. 10.7 = 9.4 8.8 Or bulla 1. 4.4 = 3.8 Aa? At) Mi1-2/ 1. 29) = 2.4 2.0 1.9 M1-1/ w. internal 3.5 = 2.4 2.7 Ped) M1/ 1. Dell — 1.6 1.4 1.2 M1/ w. 0.9 = 0.8 0.8 0.6 M2/ 1. 0.8 = 0.8 0.7 0.7 M2/ w. 0.8 = 0.7 0.6 0.6 The most recently collected specimen is AM M14166. It is an adult female snap trapped on 18 September 1979 at 1,450m on the southern slopes of Mt Sisa, Southern Highlands Province by P. Dwyer. Although most measurements are similar to those for the other M. richardsoni individuals known, it differs in a number of ways. The nasals are PROC. LINN. SOG. N.S.W., 111 (3), 1989 DY A NEW MURID FROM PAPUA NEW GUINEA longer and the mastoid width narrower than in the male specimens (Table 1). The palate does not extend as far posteriorly behind M2/ as in the Sogeri specimen. The skin of the Mt Sisa specimen is flat. It closely resembles that of the Sogeri individual except that it lacks the ginger blotching and has a small patch of pure white hairs on the throat. The most distal 25mm of the tail is pure white, while a light and dark mottled area extends a further 30mm proximally. It is unclear as yet whether the differences between AM M14166 and the other M. richardsoni specimens represent sexual dimorphism, intra, or possibly even interspecific variation. However from the specimens described here it is clear that this species, or taxa closely related to it, are relatively widely distributed across New Guinea, both geographically and in differing environments. CONCLUSION Microhydromys musseri n. sp. is described. Known from a single specimen collected in mossy forest high in the Torricelli Mountains, it differs in many aspects of its mor- phology from M. richardson. Three additional specimens of M. richardsoni are described. They indicate that this species has a wide geographic distribution and that it exhibits considerable morphological variation.’ Microhydromys richardsont can inhabit drier habitats than had previously been suspected. ACKNOWLEDGEMENTS In particular I would like to thank Dr Guy Musser of the American Museum of Natural History for his invaluable comments upon an earlier draft of this paper, and for supplying details of the Microhydromys specimens held in the American Museum. Mr Ken Aplin of the University of New South Wales also kindly commented upon this manuscript and made many useful suggestions. I would also like to thank Dr Allan Allison of the Bishop Museum for making the hydromyine specimens held in the Bishop Museum, Hawaii, available for me to study, also Mr Alan Ziegler for his help in this matter. Ms Tish Ennis produced the figures. References MENZIES, J. and DENNIS, E., 1979. — Handbook of New Guinea Rodents. Wau Ecology Institute Handbook No. 6. MISONNE, X., 1969. — African and Indo-Australian muridae. Evolutionary trends. Annis Mus. r. Afr. cent. 172: 1-219. MUSSER, G., 1981. — The giant rat of Flores and its relatives east of Borneo and Bali. Bull. Amer. Mus. Hist. 169: 69-175. MUSSER, G., 1982. — Results of the Archbold Expeditions. No. 110. Crunomys and the small-bodied shrew rats native to the Philippine Islands and Sulawesi (Celebes). Bull. Amer. Mus. Nat. Hist. 174: 1-95. OsGooD, W. H., 1945. — A new rodent from Dutch New Guinea. Fieldiana (Zoology) 31: 1-2. SMITHE, F. B., 1974. — Naturalist’s Colour Guide Supplement. New York: American Museum of Natural History. Tate, G. H. H., 1951. — Results of the Archbold Expeditions. No. 65. The rodents of Australia and New Guinea. Bull. Amer. Mus. Nat. Hist. 97: 183-430. Tare, G. H. H. and ARCHBOLD, R., 1941. — Results of the Archbold Expeditions. No. 31. New rodents and marsupials from New Guinea. Amer. Mus. Novit. 1101. THOMAS, O., 1898. — On the mammals obtained by Mr John Whitehead during his recent expedition to the Philippines. Trans. Zool. Soc. London 14: 377-412. ZIEGLER A. C., 1981. — Petaurus abidi, a new species of glider (Marsupialia, Petauridae) from Papua New Guinea. Aust. Mammal. 4: 81-88. PROG. LINN. SOG. N.S.W., 111 (3), 1989 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 111 NUMBER 4 Tidal and Diel Variations in the Abundance of Larval Fishes in Botany Bay, New South Wales, with Emphasis on Larval Silverbiddy Gerres ovatus (Fam. Gerreidae) and Gobies (Fam. Gobiidae) ALDO S. STEFFE STEFFE, A. S. Tidal and diel variations in the abundance of larval fishes in Botany Bay, New South Walcs, with emphasis on larval silverbiddy Gerres ovatus (Fam. Gerreidac) and gobies (Fam. Gobiidae). Proc. Linn. Soc. N.S.W. 111 (4), 1989: 225-232. Surface plankton samples were collected from an area of strong tidal flow in Botany Bay, during early autumn 1981, to examine tidal and diel variations in the abundance of larval fishes. The sampling program was restricted to one 24 hour period, and yielded 2,898 larvae consisting of 30 distinct larval types from 21 families. Larval gobies (Fam. Gobiidae) and silverbiddy Gerres ovatus (Fam. Gerreidae) dominated the assemblage, and together accounted for 82.5% of the standardized total catch. On the selected sampling day the composition of the larval assemblage differed at cither end of the tidal range. Gobiid larvae were more abundant at low tide both day and night, whereas abundances of G. ovatus were greater at high tide both day and night. Night catches of gobiid and G. ovatus larvae were greater than day catches. The great majority of G. ovatus larvae had deflated gas bladders during the day and inflated gas bladders at night. The limitations of these findings are acknowledged; due to the short sampling period it is possible that effects of other variables were confounded with effects of the main factors being tested. Aldo S. Steffe, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109; manuscript recewed 15 March 1989, accepted for publication 19 July 1989. INTRODUCTION Although the nursery function of Australian estuaries has been studied (Lenanton, 1977; Robertson, 1980; SPCC, 1981; Young, 1981; Bell et al., 1984; Middleton et al., 1984) little is known about larval fish distributions or the factors which affect larval recruit- ment in these estuarine areas (Steffe and Pease, 1988). The common problem facing both recruiting larvae and those already in the estuary is how to maintain their position in the estuary and avoid being flushed out on the ebb tide. Weinstein et al. (1980) proposed that successful recruitment and/or retention in a stratified estuarine system involved selective vertical migrations by larval fishes in conjunction with changes in tide and photoperiod. There is also evidence that the larvae of other estuarine dependent species respond to tidal or diel stimuli, or both (Fore and Baxter, 1972; Graham, 1972; Eldridge, 1977; Melville-Smith et al., 1981; Fortier and Leggett, 1982; Norcross and Shaw, 1984; Roper, 1986), but the mechan- ism(s) by which larval fishes achieve this are poorly understood. Here, the findings of a sampling program designed to analyse variation in larval fish abundances with respect to changes in tidal and diel conditions are reported. The main question asked was: on the selected sampling day did larval fish abundances at the surface vary with respect to changes in tide and/or diel conditions? MATERIALS AND METHODS Study Area Botany Bay (34°01’S, 151°11’E) is a large, semi-landlocked estuary on the east coast of Australia (Fig. 1). It is dominated by ocean swell and wind waves (Roy e¢ al., PROG. LINN. SOC. N.S.W., 111 (4), 1989 226 LARVAL FISHES IN BOTANY BAY 1980), is vertically well mixed (Rochford, 1951), and at most times is best described as a marine embayment. Fig. 1. Map of Botany Bay showing location of sampling station. Abbrev.: TP = Towra Point. A sampling station (Fig. 1) was selected off Towra Point because of its position in the main tidal stream and its close proximity to large Poszdonia australis and Zostera capricorn’ seagrass meadows, and an extensive mangrove stand. These habitats are important fish nursery areas (Bell e¢ al., 1984; Middleton et al., 1984). Samples were collected during March (early Autumn) as juveniles of many economically valuable fishes are most abundant in the Bay shortly after this period (Bell, 1980; SPCC, 1981; Bell et al., 1984; Middleton et al., 1984). The maximum depth at the sampling station was about 4m. Field Procedures The sampling program was designed to analyse variances in larval fish numbers between stage of tide (high vs low) and diel condition (day vs night) at one station. The design was orthogonal and required that both high and low tides be sampled during day- light, and at night. High and low tide were defined as a two hour period with the tidal prediction at its centre. Four replicate samples were collected at each consecutive high and low tide over the 24 hour sampling period. Tidal height ranged from 0.2-1.6 m above I.S.L.W. during the sampling period. Day sampling commenced 40 minutes after sunrise (05.57h) on 19 March 1981 and PROC. LINN. SOC. N.S.W., 111 (4), 1989 A. S. STEFFE Dah night sampling commenced one hour after sunset (18.09h). Plankton samples were collected from a small (4m) boat using a net with a square mouth (area 0.25m?’), a mesh of 500 wm, and a length of about 2.5m. The net was towed in a circular path at the surface at a constant speed of about 2m sec? for 5 minutes (+ 5 sec.). This procedure kept the net out of the engine wash and thus avoided unnecessarily increasing net escapement. A General Oceanics (model 2030) flowmeter was used to measure the volume of water filtered. Volume filtered per tow averaged 133.7m/? (s.d. 13.2). A salinity (+ 0.1°/0c) and water temperature (+ 0.2°C) reading was taken during each of the consecutive tides sampled. Samples were preserved immediately after collection in 5% Steedman’s preser- vative and were sorted entirely under a dissecting microscope. All larval fishes were identified to the lowest possible taxonomic level and then stored in 4% buffered forma- lin. Standardized larval abundances are expressed as the number of larvae per 100m? water filtered. Terminology follows the definitions of Leis and Rennis (1983). Size Distribution and Gas Bladder Inflation Incidence Some 200 day-caught and 200 night-caught larvae of Gerres ovatus and of gobiids were randomly selected. These larvae were measured to the nearest 0.1 mm using a dissecting microscope with mounted ocular micrometer. Notochord length was recorded for preflexion and flexion stages whilst standard length was measured for post- flexion stages. The numbers of larvae with deflated, or inflated gas bladders were then recorded separately for day-caught and night-caught G. ovatus only. This was easily done as the gas bladder in this species is clearly visible when inflated. This procedure was not repeated for gobiid larvae as they have a permanently inflated gas bladder. RESULTS The sampling program yielded 2,898 larval fishes consisting of 30 distinct larval types from 21 families (Table 1). Silverbiddy Gerres ovatus and gobiid larvae dominated the larval fish assemblage. G. ovatus larvae accounted for 31.3% of the standardized total catch whilst gobiids (four spp.) made up 51.2% of this total. The remaining 19 families contributed 17.5% of the standardized total catch but were not sufficiently abundant, or were present in too few samples, to warrant statistical analysis (Table 1). More Gerres ovatus were caught at night and during high tide (Fig. 2a, Table 2). The interaction term was not significant. Similarly, significantly more gobiid larvae were caught at night, however unlike Gerres ovatus, significantly more gobiids were caught during low tide (Fig. 2b, Table 2). There was no significant interaction (Table 2). Total fish larvae reflected the contrasting patterns of Gerres ovatus and gobiids. Night catches were significantly greater than day catches, but there was no significant tidal effect or interaction (Fig. 2c, Table 2). The length frequency of night-caught Gerres ovatus larvae was similar to that for day caught larvae (Kolmogorov-Smirnov test, Dmax. = 0.075, p> >>0.05) (Fig. 3a). Ninety-eight percent of day-caught larvae (n = 200) were found to have deflated gas bladders and ninety-five percent of night-caught larvae (n = 200) had strongly inflated gas bladders. The length frequency distribution of gobiid larvae caught during daylight was significantly different to that of gobiids caught at night (Kolmogorov-Smirnov test, Dmax. = 0.23, p<0.001) (Fig. 3b). In contrast to G. ovatus, more larger gobiid larvae were taken at night. Salinity and water temperature did not fluctuate greatly during sampling (temp. PROC. LINN. SOC. N.S.W., 111 (4), 1989 228 LARVAL FISHES IN BOTANY BAY 22.8-24.0°C; sal. 32.7-34.0°/o0) and rainfall had not been reported for the previous ten days in the area (Bureau of Meteorology, 1981). TABLE 1 Number of distinct larval types, occurrence, and the percentage of the standardized total catch for each taxon in the larval assemblage. Note that each larval type may not be monospecific No. Larval Occurrence % Standardized ‘Taxon ‘Types (Max. = 16) total catch Gobiidac 4 16 51.2 Gerreidac | 16 31.3 Ambassidae 2 15 4.7 Syngnathidac 4 16 3.7 Blenniidae 2 12 1.5 Sillaginidae 1 8 0.7 Sparidac 2 7 0.7 Monacanthidac 1 7 0.5 Carangidac | 5 0.4 Atherinidac 1 4 0.3 Anguilliformes 1 3 0.2 Clupeidac 1 5 0.2 Hemirhamphidac 1 1 0.1 Platycephalidae 1 1 0.1 Pempherididac 1 3 0.1 Soleidac 1 3 0.1 Tetraodontidac 1 3 0.1 Scorpaenidac 1 1 Mugilidac 1 1 Sphyraenidac 1 1 0.1 Callionymidac 1 1 Damaged larvac = 16 4.0 TABLE 2 F ratios and significance levels derived from two way fixed effects ANOVA of effects of tade (high vs low) and diel period (day vs night) for Gerres ovatus, Gobiidae and sample totals. Data tested for heteroscedasticity using Cochran’ test (p< 0.1). A = log transformed data, B = raw data F ratio Source of Gerres Sample Variance df ovatus’ Gobiidac® Totals? Tide 1 16.10** 5.7 0.08 NS Dicl 1 1D. So" 19) WSS" il le TxD 1 1.85 NS 0.06 NS 0.08 NS Residual 12 *» <0.05, ** p <0.01, *** p <0.001, NS p >0.05 DISCUSSION Gerres ovatus larvae were significantly more abundant on high tides whilst gobuid larvae were caught in significantly greater numbers during low tides. As G. ovatus and numerous gobiid species are known to spawn within Botany Bay (State Pollution Control Commission, 1981) their centres of larval abundance may have been expected to coincide. Yet, they have become spatially separated, occurring at either end of the tidal range. This larval distribution was found both during daylight and at night. Two hypotheses, not mutually exclusive, to explain this are: (1) spawning aggregations of G. PROC. LINN. SOC. N.S.W., 111 (4), 1989 (a) 100 5 S) 80 ti a 60 a ra = 40 — ol = 20 NIGHT HIGH TIDAL STAGE (c) N@ LARVAE PER 100m3 250 200 150 100 50 AGS) SBE EE 229 160 140 wae NIGHT 100 80 60 40 NO LARVAE PER 100m? 20 HIGH LOW TIDAL STAGE NIGHT DAY HIGH LOW TIDAL STAGE Fig. 2. Mean and + one SE for the standardized larval abundances at each of the consecutive tides, both day and night, over the 24 hour sampling period for (a) Gerves ovatus, (b) Gobiidae, and (c) sample totals. PROG. LINN. SOC. N.S.W., 111 (4), 1989 230 LARVAL FISHES IN BOTANY BAY (a) (b) 60 50 40 pce eew em eww ew een’ 30 FREQUENCY FREQUENCY 20 Bosooosoo00K 10 7 DTS Ran eS IGS 7 123. > 42 35006 LENGTH (mm) LENGTH (mm) Fig. 3. Length frequencies of randomly selected day-caught (n = 200) and night-caught (n = 200) larvae for (a) Gerres ovatus, and (b) Gobiidae. Solid lines denote day-caught, and broken lines denote night-caught. ovatus occur near the Bay entrance and seaward of gobiid spawning sites; and (2) both groups spawn at a similar distance into the Bay, but the demersal eggs of gobiids are more effective than the pelagic eggs of G. ovatus at reducing the net transport of offspring seaward from spawning sites by currents. This could occur because demersal eggs, unlike pelagic eggs, are not subject to passive transportation by currents, and because at hatching, larvae from demersal eggs tend to be relatively larger, more developed, with better swimming capabilities than those from pelagic eggs (Steffe and Pease, 1988). Larval catches of Gerres ovatus and gobiids were found to be significantly greater at night (Table 2). There was no difference in the size structure of G. ovatus between day and night (Fig. 3a) suggesting that differential net avoidance was minimal and that the higher night catches could be mainly attributed to the effects of diel vertical migration. PROC. LINN. SOC. N.S.W., 111 (4), 1989 AS. SUEFFE 231 The effects of differential net avoidance and diel vertical migration could not be separated for gobiids as more large larvae were caught at night (Fig. 3b). Most Gerres ovatus larvae had deflated gas bladders during the day whilst the reverse was true at night. This phenomenon appears to be common and occurs in many taxo- nomically diverse teleost groups (Hunter and Sanchez, 1976; Leis and Rennis, 1983; Liew, 1983; Hoss and Phonlor, 1984; Kitajima et al., 1985; A. Steffe, unpub. data). Hunter and Sanchez (1976) found that nocturnal gas bladder inflation can provide considerable energy savings to northern anchovy Engraulis mordax larvae by retarding sinking. It is likely that larval G. ovatus gain a similar benefit. The silverbiddy, G. ovatus, has been found to recruit almost exclusively to mangrove areas in Botany Bay (State Pollution Control Commission, 1981). The observed larval distribution of G. ovatus at a site near its preferred nursery habitat supports the hypothe- sis that larval silverbiddy may be using flood tides to assist their transportation into mangrove areas. This hypothesis appears tenable because mangrove areas are wholly dependent on tide for water exchange. The data presented and interpreted here are based on collections made during a single 24 hour period. Consequently, it is possible that the effects of other variables which were not specifically tested may have, by chance, been confounded with the effects of the tidal and diel factors examined during the selected sampling day; in view of the relatively constant temperature and salinity conditions it is unlikely that larval catches were influenced by these parameters. Further sampling, that is, the same experiment repeated on other ‘replicate’ days, is required to eliminate this possible source of error and to possibly allow the conclusions drawn here to be accepted with greater confidence. ACKNOWLEDGEMENTS I thank B. Griffiths, B. Hodgson, J. Leis, J. MacIntyre, A. Mazanov, A. Miskiewicz and M. Westoby all of whom contributed to the compilation and completion of this work. Special thanks to my parents and wife for financial support and field assistance. L. Beckley, J. Bell, W. Gladstone, J. Leis, K. McGuiness, and M. Westoby made useful suggestions which improved the manuscript. References BELL, J. D., 1980. — Aspects of the ecology of fourteen economically important fish species in Botany Bay, New South Wales, with special emphasis on habitat utilization and discussion of the effects of man- induced habitat changes. North Ryde, N.S.W.: Macquarie University, M.Sc. thesis, unpubl. — , POLLARD, D. A., BURCHMORE, J. J., PEASE, B. C., and MIDDLETON, M. J., 1984. — Structure ofa fish community in a temperate tidal mangrove creek in Botany Bay, New South Wales. Aust. J. Mar. Freshw. Res. 35: 33-46. BUREAU OF METEOROLOGY, 1981. — Monthly weather review. New South Wales. Dept. of Science, Canberra. ELDRIDGE, M. B., 1977. — Factors influencing distribution of fish eggs and larvae over eight 24 hour samplings in Richardson Bay, California. Calif. Fish and Game 63: 101-116. Fore, P. L., and BAXTER, K. N., 1972. — Diel fluctuations in the catch of larval gulf menhaden, Brevoortia patronus, at Galveston Entrance, ‘lexas. Trans. Amer. Fish. Soc. 101: 729-732. Fortier, L., and LEGGETT, W. C., 1982. — Fickian transport and the dispersal of fish larvae in estuaries. Canadian J. Fish. Aqu. Sci. 39: 1150-1163. GRAHAM, J. J., 1972. — Retention of larval herring within the Sheepscot estuary of Maine. U.S. Fish. Bull. 70: 299-305. Hoss, D. E., and PHONLOR, G., 1984. — Field and laboratory observations on diurnal swim bladder inflation-deflation in larvae of gulf menhaden, Brevoortza patronus. U.S. Fish. Bull. 82: 513-517. HUNTER, J. R., and SANCHEZ, C., 1976. — Diel changes in swim-bladder inflation of the larvae of the northern anchovy, Engraulis mordax Girard. U.S. Fish. Bull. 74: 847-855. KiTajIMA, C., TSUKASHIMA, Y., and TANAKA, M., 1985 — The voluminal changes of swim bladder of larval red sea bream Pagrus major. Bull. Japan. Soc. Sci. Fish. 51: 759-764. PROC. LINN. SOG. N.S.W., 111 (4), 1989 DBD LARVAL FISHES IN BOTANY BAY Leis, J. M., and RENNIS, D. S., 1983. — The larvae of Indo-Pacific coral reef fishes. Kensington, N.S.W.: Univer- sity N.S.W. Press and Honolulu: University of Hawaii Press. LENANTON, R. C. J., 1977. — Aspects of the ecology of fish and commercial crustaceans of the Blackwood River estuary, Western Australia. Fish. Res. Bull. WA. 19: 1-72. Liew, H. C., 1983. — Studies on flatfish larvae (Fam. Psettodidae and Bothidae, Pleuronectiformes) in the shelf waters of the central Great Barrier Reef, Australia. Townsville, Qld.: James Cook University, M.Sc. thesis, unpubl. MELVILLE-SMITH, R., BAIRD, D., and WOOLDRIDGE, T., 1981. — The utilization of tidal currents by the larvae of an estuarine fish. Sth. African J. Zool. 16: 10-13. MIDDLETON, M. J., BELL, J. D., BURCHMORE, J. J., POLLARD, D. A., and PEASE, B. C., 1984. — Structural differences in the fish communities of Zostera capricornt and Posidonia australis seagrass meadows in Botany Bay, New South Wales. Aquat. Bot. 18: 89-109. Norcross, B. L., and SHAW, R. F., 1984. — Oceanic and estuarine transport of fish eggs and larvae: a review. Trans. Amer. Fish. Soc. 113: 153-165. ROBERTSON, A. I., 1980. — The structure and organization of an eelgrass fish fauna. Oecologia 47: 76-82. ROCHFORD, D. J., 1951. — Studies in Australian estuarine hydrology. I. Introductory and comparative features. Aust. J. Mar. Freshw. Res. 2: 1-116. Roper, D. S., 1986. — Occurrence and recruitment of fish larvae in a northern New Zealand estuary. Est. Coast. Shelf Scr. 22: 705-717. Roy, P. S., THoM, B. G., and WRIGHT, L. D., 1980. — Holocene sequences on an embayed high-energy coast: an evolutionary model. Sedimentary Geology 26: 1-19. STATE POLLUTION CONTROL COMMISSION, 1981. — The ecology of fish in Botany Bay — biology of commer- cially and recreationally valuable species. SPCC Rep. No. BBS 23B, 1-287. STEFFE, A. S., and PEASE, B. C., 1988. — Diurnal survey of ichthyoplankton abundance, distribution and seasonality in Botany Bay, New South Wales. Proc. Linn. Soc. N.S.W. 110: 1-10. WEINSTEIN, M. P., WEISS, S. L., HODSON, R. G., and GERRY, L. R., 1980. — Retention of three taxa of post- larval fishes in an intensively flushed tidal estuary, Cape Fear River, North Carolina. U.S. Fish. Bull. 78: 419-436. YOUNG, P. C., 1981. — Temporal changes in the vagile epibenthic fauna of two seagrass meadows (Zostera capricornt and Posidonza australis). Mar. Ecol. Prog. Ser. 5: 91-102. PROC. LINN. SOC. N.S.W., 111 (4), 1989 Redescription of Austrochthonius australis Hoff (Chthonidae: Pseudoscorpionida) CLARICE M. A. KENNEDY KENNEDY, C. M. A. Redescription of Austrochthonius australis Hoff (Chthoniidae: Pseudoscorpionida). Proc. Linn. Soc. N.S.W. 111 (4), 1989: 233-240. An examination of approximately 150 specimens of Austrochthonius australis Hoff from various locations in southeastern Australia suggests a considerable size variation within the species when compared with the two published descriptions of adults (Hoff, 1951; Beier, 1966). For the first time the nymphal stages are described. Clarice M. A. Kennedy, School of Biological Sciences, Macquarie University, North Ryde, Aus- tralia, 2109, manuscript received 7 February 1989, accepted for publication 19 July 1989. INTRODUCTION A. australis 1s widely distributed in a variety of habitats throughout Australia. The description of the holotype is based on a single female specimen from Mt Slide, | Victoria. Beier (1966) has briefly described male and female specimens based on an examination of material from Western Australia, Victoria, New South Wales, Australian Capital Territory and Tasmania. Investigation of specimens from Western Australia, identified as A. australis, has revealed a number of differences from the holotype suggesting that an undescribed species is represented. In addition, individuals of a population in the Sydney region are much smaller, when compared with the two published descriptions of the adult by Hoff (1951) and Beier (1966); this prompted examination of some 150 specimens from various locations in Victoria, New South Wales and Tasmania to determine if specimens identi- fied as A. australis are in fact a single species. Apart from size, specimens examined are morphologically similar to the holotype. The results of this research are presented below and represent a more compre- hensive description of the species which for the first time, incorporates the nymphal stages. MATERIALS AND METHODS Specimens were borrowed from the following institutions: American Museum of Natural History, New York; Australian Museum, Sydney; Museum of Victoria, Melbourne and 34 samples from the private collection of Dr. M. S. Harvey, Western Australian Museum, Perth. Measurements are based on the examination of at least 5-15 adults of each sex from various locations in south eastern Australia made in accordance with those described by Chamberlin (1931). Figures in parentheses are female values and follow those of the male. The morphological nymphal information is based on measurements of 10 specimens of each stage collected from the Sydney region. Nymphs from the other regions within the distribution of the species are, at present, unavailable for comparison. Abbreviations for chelal trichobothria and setal formulae follow those devised by Chamberlin (1931). Genitalia terminology follows that of Legg (1975). Specimens of A. australis from the Sydney region are deposited in the Australian Museum, 6 © (KS 20186), 6 9 (KS 20187), 6 nymphs (KS 20188). PROG. LINN. SOC. N.S.W,, 111 (4), 1989 234 REDESCRIPTION OF PSEUDOSCOR PION SYSTEMATIC DESCRIPTION CHTHONIIDAE Hansen 1893 Austrochthonius Chamberlin 1929 Type species: Chthonius chilensis Chamberlin 1923 Austrochthonius australis Hotf (Figs 1-4) Diagnosis: Chelicera with 5 setae on hand; palpal teeth quadrate and separated; 6 pin- nate coxal spines present on coxa 2; intercoxal tubercle absent. L/W ratio of chela (with pedicel) range 1:1.05-1.33 (1: 1.03-1.45) x longer than broad. Description: Adults. Colour light golden brown; carapace surface smooth; pleural membrane with series of irregularly spaced longitudinal ridges, each with minute, elevated, white nodules (Fig. 1A). Pedipalp: trochanter stout, femur with lateral margins virtually parallel, abruptly converging distally, L/W ratio range 1: 1.00-1.65 (1: Wa VOam . ; g . | VOmpm Fig. 1. Austrochthonius australis Hoff, scanning electron micrographs, male: A, pleural membrane; B, chelicerac, dorsal aspect; C, right cheliccra movable finger, ventral aspect; D, left chelicera, serrula exterior proximal blades; E, right chelicera, scrrula exterior distal blades, ventral aspect; F, right chelicera, scrrula exterior distal blades, dorsal aspect. PROC. LINN. SOG. N.S.W., 111 (4), 1989 C. M. A. KENNEDY D335) 1.00-1.65), tibia calyciform; chelal hand broad and shorter than chelal fingers, lateral margins very slightly rounded, poorly developed pedicel, L/W ratio of chela (with pedicel) range 1: 1.05-1.33 (1: 1.03-1.45), chela (without pedicel) 1: 1.03-1.31 (1: 1.01-1.42) x longer than broad. Chelal fingers long and tapering, fixed finger with 8 trichobothria, 26 and zsh medial on dorsum of chelal hand; movable chelal finger with 4 trichobothria; marginal teeth quadrate, separated and retrorse on fixed finger, teeth smaller on movable finger becoming broader with rounded apices and laterally fused proximally, 46-50 (0) 45-46 (9 ) (Fig. 4T); venom apparatus absent. Chelicera large, stout with 5 setae on hand, fingers cross distally (Figs 1B, 4S), fixed finger with reticulate sculpture, teeth 15-16 (0”) 15-17 (Q ), distal one large, remainder diminishing in size proximally; movable finger with 18 well-spaced smaller teeth which become almost indistinguish- able proximally; lamina exterior absent. Serrula exterior detached in upper half with 14- Fig. 2. Austrochthonius australis Hoff, scanning electron micrographs, male: G, right chelicera, flagellum blades, ventral aspect; H, female, right chelicera showing galea a sclerotic tubercle; I, carapace; J, male, epistome; K, coxal spines; L, enlargement of pinnate coxal spine blades. 15 (0°) 14(Q ) lamellae (Fig. 1C), lamellae relatively broad, elongate, with parallel sides proximally, terminally each blade digitate (Fig. 1D) becoming broad distally (Figs 1E, PROC. LINN. SOC. N.S.W., 111 (4), 1989 236 REDESCRIPTION OF PSEUDOSCORPION Fig. 3. Austrochthonius australis Hoff, scanning electron micrographs: M, male genital region, external aspect; N, female genital region, external aspect; P, male genitalia, internal aspect; Q, female genitalia, internal aspect. F); galea a sclerotic tubercle in 9 (Fig. 2H), absent in ©’; galea seta basad of mid-point of movable finger. Flagellum with 8 stalk-like blades each arising separately along a diagonal slit basad of fixed cheliceral finger (Fig. 1C) becoming unipinnate distally (Fig. 2G). Carapace: subquadrate, surface smooth; anterior margin slightly serrate laterally with prominent, blunt, dentate epistome (©’) (Fig. 2J), more or less acute (@ ), weakly reticulate sculpture; lateral margins distinctly narrowed posteriorly (Fig. 21); posterior margin relatively straight; setae long, acuminate 4: 2: 12 (0), 4: 2: 12-14 (2 ); L/W ratio PROG. LINN. SOG. N.S.W., 111 (4), 1989 Cc. M. A. KENNEDY Zh (based on ocular width) range 1: 1.00-1.13 (1: 1.03-1.36), (based on posterior width) 1: 1.07-1.52 (1: 1.12-1.33) x longer than broad. Anterior eye well developed, posterior eye flattened disc. Tergal chaetotaxy: setae relatively long, acuminate, O 4: 4: 3-4: 2-4: 6: Goiioa a Oa) Oa Ae ee O07 014-0: 2. UinIsenlate. Goxalichaetotaxya 6 pinnate coxal spines placed anterior-ventrally along proximal half of coxa 2 (Figs 2K, L) Oe eae srl O a DO 23-45 One 4) 2:92 1-2)02.3 2. Ons nO: 2-32.45. 102 2213) Intercoxal tubercle absent. Male genitalia (Fig. 3P): ejaculatory canal atrium (ejca) pear-shaped; roof of each medial diverticulum thickened (trmd) to form two elongated structures, anterior horns of each free but fused posteriorly, in mid-line is atrium of posterior dorsal gland (apdg). Cuticle of genital atrium thickened and extended to form apodemes; dorsal apodeme (da) well developed with Y-shaped structure below ejacu- latory canal atrium (ejca), posterior extension of Y forms anterior crest of dorsal apodeme (acda), posterior to atrium of posterior dorsal gland (apdg) the posterior crest of the dorsal apodeme (pcda) arises in mid-line above duct of median genital sac; lateral apodemes (la) reduced and thickened to provide attachment for muscles of ejaculatory canal and support for anterior region of genital region of genital atrium. Chaetotaxy: anterior operculum with 8-9 relatively long setae along anterior margin, 4-5 along posterior margin which is deeply concave medially; posterior operculum with 8 relatively long acuminate setae on each side of a deep median notch. These setae project into the centre forming a grill-like pattern across the genital atrium (Fig. 3M). Female genitalia (Fig. 3Q): lateral cribriform plates (Icp) clearly differentiated with numerous pores; median cribriform plate (mcp) — pores scattered over the surface with some aggregations. Chaetotaxy: anterior operculum a triangular plate; posterior operculum a simple elongate plate (Fig. 3N). Sternites with reticulate sculpture, chaetotaxy: O 0: 13- dee Wa /2 dee Gee Saas Ge (Oe (OR 4-153 Ges 23 © (Os ile We os (Os 403 (3 Os Os Oe A, Seizie Nora. acuminate. Sternites 4-11 uniseriate. Dimensions (mm): body length 0.75-117 (0.77-1.62); pedipalp: femur 0.21- 0.36/0.07-0.10 (0.21-0.46/0.07-0.10), chela (with pedicel) 0.43-0.61/0.09-0.13 (0.45- 0.70/0.10-0.16), chela (without pedicel) 0.41-0.59/0.09-0.13 (0.41-0.68/0.10-0.16), movable finger length 0.25-0.37 (0.28-0.45). Carapace 0.27-0.37/ocular width 0.27- 0.41/posterior width 0.21-0.34 (0.28-0.46/ocular width 0.30-0.48/posterior width 0.26-0.43). Tritonymph. L/W ratio: pedipalpal trochanter 1.00-2.00, femur 2.28-3.00, tibia 1.28-1.42, chela (with pedicel) 3.55-4.33, chela (without pedicel) 3.33-4.11 x longer than broad. Fixed finger with 7 trichobothria, movable finger with 3 trichobothria, zsb and sb absent; serrula exterior of chelicera with 11-12 broad, elongate lamellae, each blade digitate terminally. Carapace: anterior margin slightly serrate laterally, epistomal process prominent; posterior margin relatively straight; lateral margin narrowing at point of union with abdominal segments, 4: 2: 8, setae long, acuminate. L/W ratio (based on ocular width) 1.08-1.25, (based on posterior width) 0.96-1.00 x longer than broad. Tergal chaetotaxy: 4: 4: 4: 4: 4-6: 4-6: 6: 6: 6: 4: 4-6: 2. Sternal chaetotaxy: 0: 3-4: 6-7: 6: 6: 4: 4: 6: 6: 6-7: 5-6: 2. Coxal chaetotaxy: 4-5 coxal spines in medial position Alone antcHonimaeimon2nd coxa, 22) 11023010: 302-35 Onl 2-45 Olena Dimensions (mm): body length 0.66-0.79; pedipalp: trochanter 0.07-0.09/0.05- 0.07, femur 0.16-0.21/0.07, tibia 0.09-0.10/0.07, chela (with pedicel) 0.32-0.39/0.07-0.09, chela (without pedicel) 0.30-0.37/0.07-0.09, movable finger length 0.19-0.27; carapace 0.27-0.30/ocular width 0.27-0.31/posterior width 0.22-0.25. Deutonymph. L/W ratio: pedipalpal trochanter 1.20-1.40, femur 2.00-3.20, tibia 1.00-1.75, chela (with pedicel) 3.75-4.50, chela (without pedicel) 3.28-4.16 x longer than broad. Fixed finger with 6 trichobothria, movable finger with 2 trichobothria, zsd, esd, sb, 6 absent; serrula exterior of chelicera with 9-10 broad, elongate lamellae, each blade PROC. LINN. SOC. N.S.W., 111 (4), 1989 238 REDESCRIPTION OF PSEUDOSCORPION ee! 0.14¢m Fig. 4. Austrochthonius australis Hoff, male: S, right chelicera, dorsal aspect; T, right chela, lateral aspect; Ta, inset, enlargement showing dentition type (not to scale); U, first right leg; V, fourth right leg; W, left pedipalp, dorsal aspect. PROC. LINN. SOC. N.S.W., 111 (4), 1989 Cc. M. A. KENNEDY 239 digitate terminally. Carapace: anterior margin serrate, epistome prominent; 4: 2: 8: 3-6 relatively long setae, 1.05-1.18 x longer than broad (ocular and posterior width equal). Tergal chaetotaxy: 4: 4: 4: 2-4: 4-6: 4-6: 4-5: 5-6: 4-5: 4-6: 5-6: 2, setae, long, acuminate posteriorly. Sternal chaetotaxy: 0: 4: 4: 4-6: 4-6: 6: 4: 4: 4-6: 4-6: 6: 2. Sternites 3-5 divided. Coxal chaetotaxy: 3-4 coxal spines in medial position along anterior margin of ZnCl, COs 22 Ze i, Ws Zevs OO Zeal, Os evs a, Oe ale Dimensions (mm): body length 0.48-0.63; pedipalp: trochanter 0.06-0.07/0.05, femur 0.12-0.16/0.04-0.07, tibia 0.07/0.05-0.07, chela (with pedicel) 0.25-0.28/0.06-0.07, chela (without pedicel) 0.23-0.27/0.06-0.07, movable finger length 0.14-0.19; carapace 0.19-0.23/0.16-0.20. Protonymph. L/W ratio: pedipalpal trochanter 1.25-1.33, femur 2.20-3.00, tibia 1.25-1.50, chela (with pedicel) 3.33-4.00, chela (without pedicel) 3.16-3.80 x longer than broad. Fixed finger with 3 trichobothria, ist, et, eb present; movable finger with 1 trichobothrium, ¢ present; serrula exterior of chelicera with 7-8 broad, elongate lamellae. Carapace: anterior margin smooth, slightly dentate medially, 4: 2: 5: 3 setae, 1.06- 1.16 x longer than broad (ocular and posterior width equal). Tergal chaetotaxy: 2: 2-3: 2- 3; 2-3: 42 3: 3-4; 3: 4: 3-42 4-5: 2. Sternal chaetotaxy: 0; 2-323: 4: 2-3: 2-4: 4: 4: 3-4: 3-4: 4: 2. Coxal chaetotaxy: 3-4 small coxal spines along anterior margin of coxa 2, 2: 2: 1, 0: 2 (0. Os 2 ©, Os WE7s sips deal Dimensions (mm): body length 0.36-0.45; pedipalp: trochanter 0.04-0.05/0.03- 0.04, femur 0.09-0.11/0.03-0.05, tibia 0.05-0.06/0.04, chela (with pedicel) 0.17-0.22/0.05- 0.06, chela (without pedicel) 0.16-0.21/0.05-0.06, movable finger length 0.11-0.14; carapace 0.14-0.17/0.12-0.16. DISCUSSION Only two species of the genus Austrochthonius Chamberlin are recorded from Australia namely: A. australis Hoff and A. cavicola Beier (1968). Results of this investiga- tion are based on the examination of specimens from locations ranging from lat. 33°45'S, long. 150°57’E to lat., 41°49’S, long. 145°37’E and suggest a considerable size variation exists within A. australis which is not consistent with clinal variation. While size is important, body length was not considered a reliable guide in itself because measurements can vary considerably due to a number of factors; for example, the freshness of material, the inclusion of gravid specimens, the nature of preservative and length of time in fixative. Consequently, the results of this research are based on measurements of important specific morphological features which remain relatively constant within populations. A comprehensive description of the nymphal population is presented for the first time and is therefore applicable only to the small specimens recovered from the Cumberland State Forest, Sydney, N.SW. Lack of material representative of southeastern Australia precluded a reliable assessment in this respect. ACKNOWLEDGEMENTS I wish to thank Dr. N. I. Platnick (American Museum of Natural History), Dr. M. Gray (Australian Museum), Dr. M. S. Harvey (Western Australian Museum) for loan of material and Dr. N. N. Tait for reviewing the manuscript. References BEIER, M., 1966 — On the Pseudoscorpionidea of Australia. Aust. J. Zool., 14: 265-283. ——, 1968 — Some cave dwelling Pseudoscorpionidea from Australia and New Caledonia. Rec. S. Aust. Mus., 15: 757-758. PROC. LINN. SOG. N.S.W., 111 (4), 1989 240 REDESCRIPTION OF PSEUDOSCORPION CHAMBERLIN, J. C., 1923 — On two species of pseudoscorpion from Chile with a note on one from Sumatra. Revista Chilena de Historia Natural, 27: 187-190. ——, 1929 — A synoptic classification of the false scorpions or chela-spinners, with a report on a cosmo- politan collection of the same, Part I. The Heterosphyronida (Chthoniidae) (Arachnida — Chelon- etida). Ann. Mag. nat. Hist., 4: 50-80. , 1931 — The arachnid order Chelonethida. Stanford Univ. Publs. (Biol. ), 7: 1-284. HANSEN, H. J., 1893. — Organs and characters in different orders of Arachnids. Ent. Medd., 4: 232. Horr, C. C., 1951. — New species and records of Chthoniid pseudoscorpions. Am. Mus. Novit., 1483: 1-13. LeGcG, G., 1975. — The genitalia and associated glands of five British species belonging to the family Chthoniidae (Pseudoscorpiones: Arachnida). J. Zool. London, 177: 99-121. PROC. LINN. SOC. N.S.W., 111 (4), 1989 Pied Currawongs (Strepera graculina): their Diet and Role in Weed Dispersal in suburban Sydney, New South Wales ROBIN A. BUCHANAN BUCHANAN, R. A. Pied Currawongs (Strepera graculina): their diet and role in weed dispersal in suburban Sydney, New South Wales. Proc. Linn. Soc. N.S.W. 111 (4), 1989: 241-255. In this dietary study of an abundant population of pied currawongs (Strepera graculina) 1009 regurgitated pellets, collected over 25 months, were analysed. The percentage of pellets containing plant parts, mostly seeds of fleshy fruit, was high throughout the year (79-98%). The fruit of introduced plants was present in 45-91% of pellets containing plant material. The seeds of 46 species, of which 36 were introduced, were identified in their pellets. Fruits of the family Oleaceae, including the three introduced species, Ligustrum sinense (small-leaved privet), Ligustrum lucidum (large-leaved privet) and Olea africana (wild olive) were the most significant part of the currawong diet for three months of the year, when at least one of these species was present in 54-74% of the pellets analysed. Other major plant species in the diet included the introduced Pyracantha angustifolia, Morus nigra, Ochna atropurpurea, Solanum pseudocapsicum, and the native Elaeocarpus reticulatus. The proportion of pellets containing animal parts decreased from a value of 50- 75% in the warmer months to 11-12% in the coldest months of the year. Few vertebrate remains were found in the pellets; the major animal components identified were bull- ants (Myrmecia spp.), beetles, and other insects. Robin A. Buchanan, 22 Alicia Road, Mt. Kuring-gat, Australia 2080; manuscript received 22 December 1987, accepted for publication in revised form 19 April 1989. INTRODUCTION A varied diet, intelligence and bold behaviour have contributed to making the pied currawong (Strepera graculina) one of the most successful and abundant birds in suburban environments throughout eastern Australia (Readshaw, 1968; Wimbush, 1969; Blakers et al., 1984). Within twenty years of European colonization, pied currawongs had exploited new habitats and foods provided by settlement (Currey, 1966); and by the 1920’s they were listed as one of the main dispersal agents of Opuntia spp. (Anon, 1920; 1927). More recently it has become clear that they are also agents for the dispersal of two abundant privet species (Ligustrum sinense, L. lucidum) in suburban Sydney (Walsh, 1965; Vellenga, 1966; Rose, 1973; Clyne, 1980). Despite their large size (41-51cm), abundance and role in weed dispersal, the diet of pied currawongs has not been studied in detail. Rose (1973) systematically sampled regurgitated pellets, but many other accounts are largely anecdotal. Reports of their animal prey occur in Lea and Grey (1936), Marshall (1935), Roberts (1942), Readshaw (1965), Recher (1976), and Cooper and Cooper (1981) and include many insect orders as well as worms, snails, crabs, birds, mammals and carrion. Items of fruit eaten are frequently listed in catalogues of regional birdlife (Table 1). From October 1976 to October 1978 the pellets regurgitated by pied currawongs in a garden in suburban Sydney were collected and their contents analysed. Objectives of this survey were: to record details of the larger contents of pellets, relating them to seasonal abundance of dietary components; to discuss the dietary findings in relation to the role of pied currawongs in weed dispersal in the Sydney region. PROC. LINN. SOC. N.S.W., 111 (4), 1989 242 CURRAWONGS AND WEED DISPERSAL TABLE 1 Fruits and seeds reported in previous studies and additional to those recorded from Thornleigh Identification* MONOCOTYLEDONES Poaceae *Maize DICOTYLEDONES Anacardiaceae *Schinus molle *Pepper Tree *Pepper Tree Aquifoliaceae *Holly Tree Cactaceae *Prickly Pear *Prickly Pear *Prickly Pear Loranthaceae Mistletoe Loranthus Mistletoe Mistletoe Malvaceae #Lagunaria patersonia Mimosaceae # Paraserianthes lophantha Moraceae Moreton Bay Fig Ficus macrophylla Myrtaceae #Eucalyptus caesia Philesiaceae Rosaceae *Loquat *Wild Raspberry *Rowan *English Laurel Rutaceae *Orange and Lemon Santalaceae Exocarpus stricta Solanaceae *Chillies Vitaceae Native Grape x = * = introduced to Australia. # = + N = notstated. O = observation while feeding. P = pellet S = stomach content. Author Edwards 1922 Rose 1973 Cheney 1915 Bourke 1949 Vallenga 1966 Morse 1922 Anon 1920 Anon 1927 Cleland, Maiden, Ferguson & Musson 1918 Lea and Gray 1936 Keast 1958 Rose 1973 Smith, Larkins & Pegler 1984 Smith, Larkins & Pegler 1984 Agnew 1922 Robertson 1969 Smith, Larkins & Pegler 1984 Rose 1973 Roberts 1942 Marshall 1935 Vallenga 1966 Vallenga 1966 Roberts 1942 Lea & Gray 1936 Bravery 1970 Cleland, Maiden, Ferguson & Musson 1918 indigenous to Australia but not to the Thornleigh area. SITE DESCRIPTION Type of Record + MA ©O00O 242 2 s2ZQam BAwzwzz © OO's Z QO OOOZdZ Nn The common or scientific name is listed in the same form as that given in the relevant paper. The study was carried out at Thornleigh, 19km north-west of Sydney, New South Wales; area features are illustrated in Fig. 1. Thornleigh is situated on a ridge 1-3km wide with the steep-sided valleys of the Lane Cove River to the south-east and Berowra Creek to the north-west. Both valleys are forested and the land to the west of Berowra PROC. LINN. SOC. N.S.W., 111 (4), 1989 R.A. BUCHANAN 243 Creek is rural. Most of the suburban development consists of detached dwellings in gardens well vegetated with native and introduced trees and shrubs. The percentage of suburban development within a one to three kilometre radius of the study site ranges from 50 to 67% respectively, with the remainder being natural vegetation. E D railway highway ~ built-up Collection site @ Fig. 1. Map of the environs of the collection site showing the distribution of the built-up areas and native vegetation. The majority of the natural vegetation is woodland/open-forest (Specht, 1970) and is included in the Sandstone Complex, both wet and dry sclerophyll, of Specht e¢ al., (1974). Most of the slopes are free of weeds but L. sznense and other weeds are abundant along the creek and river banks. Regurgitated pellets were collected at the junction of built-up and forested areas on the eastern side of Thornleigh. The collection site was a lawn surrounded by a dense growth of shrubs and trees; pied currawongs were attracted by bread, honey, seed, meat and water. PROG. LINN. SOG. N.S.W., 111 (4), 1989 244 CURRAWONGS AND WEED DISPERSAL METHODS Pellets Pellets were collected twice a week over an area of approximately 150 m? from October 1976 to October 1978. In total 1 009 pellets were examined; the highest number sampled in any one month was 53 and the lowest 8 (Table 2). In October 1978 some 41 pellets were analysed. TABLE 2 Number of pellets analysed in the 24 month period (October 1976-September 1978) and collected in the second year 2 48 4] 2 (Analysed) 2 119 64 24 2 (Collected) The sampled pellets were divided into three classes of coherence; coherent (collected intact or almost intact), partly coherent (collected in more than one piece), not coherent (collected completely fragmented). The lengths and widths of coherent pellets were measured and the contents of coherent and partly coherent pellets were then separated by stirring them in water and filtering. All pellets separated into individual fragments with this treatment. Only the larger fragments, identifiable at 40x magnifi- cation, were noted. Component Identification Most seeds were identified by comparison with seeds from known plants and by identification of young plants grown from seeds in the pellets. Some of the rarer seeds in the pellets were identified by the National Herbarium (Sydney). Scientific and vernacu- lar names are listed in Table 3; botanical and common names follow the National Herbarium of New South Wales. Classification of invertebrates — based on Common- wealth Scientific and Industrial Research Organization, 1970 — was only attempted to class or order and rarely to genus. Many tiny invertebrate fragments remained un- identified. Some bone fragments were identified by the Australian Museum, Sydney. The number of different components, identified to class or genus for insects, and species for plants was tallied for each pellet. The plant species most frequently present in pellets was assessed by counting the number of pellets in which a species was present in each month. Bull-ants (Myrmecia species) As the number of pellets containing bull-ants showed a strong seasonal pattern, an estimate of the activity of bull-ants was obtained by observing six nests for 5-10 minutes on one afternoon a week during the second year of the study. The largest number of ants present on the surface of the nest during this time was recorded. The average number of observed bull-ants was then calculated for each month. PROC. LINN. SOC. N.S.W., 111 (4), 1989 R. A. BUCHANAN 245 TABLE 3 Species of fruit recorded in pellets Highest frequency in any one month (7%) Classification (Common Name) 00. | 10-30 30-50 50-70 70-90 GYMNOSPERMAE Cupressaceae * Juniperus sp. (Juniper) x MONOCOTYLEDONES Arecaceae *Phoenix sp. (Palm) x Asparagaceae #0* Protasparagus aethiopicus (Asparagus Fern) Asteliaceae + Cordyline rubra Phormiaceae Dianella caerulea (Flax Lily) Smilacaceae Smilax glyciphylla (Thornless Smilax) Zingiberaceae #0* Hedychium gardnerianum (Ginger Lily) DICOTYLEDONES Anacardiaceae #0* Toxicodendron succedaneum (Rhus) x Araliaceae 0* Hedera helix (Ivy) x Polyscias sambucifolia x Cornaceae * Dendrobenthamia capitata x Ebenaceae *Diospyros kaki (Persimmon) x Elaeocarpaceae Elaeocarpus reticulatus (Blueberry Ash) x Euphorbiaceae Omalanthus populifolius (Poplar-leaved Omalanthus) x Lauraceae #0* Cinnamomum camphora (Camphor Laurel) Xx Magnoliaceae 0* Magnolia grandiflora (Evergreen Magnolia) x Malvaceae + Lagunarta patersonia (Norfolk Island Hibiscus) x Meliaceae 0+ Melia azedarach (White Cedar) x Moraceae * Ficus carica (Commercial Fig) Xx Ficus rubiginosa (Port Jackson Fig) x 0* Morus alba (Mulberry) xX Myrtaceae Eucalyptus resinifera (Red Mahogany) x + Syzygium paniculatum (Lilly Pilly) x Ochnaceae #0* Ochna atropurpurea (Ochna) x Oleaceae #0* Ligustrum lucidum (Large-leaved Privet) x #0* Ligustrum sinense (Small-leaved Privet) Xx #0* Olea africana (Wild Olive) Xx Phytolaccaceae #0* Phytolacca octandra (Ink Weed) x Pittosporaceae Pittosporum undulatum (Pittosporum) x x we Km KK PROC. LINN. SOC. N.S.W., 111 (4), 1989 246 CURRAWONGS AND WEED DISPERSAL TABLE 3 (Cont’d.) Highest frequency in any one month (%) Classification (Common Name) Proteaceae Persoonia pinifolia (Pine-leaf Geebung) Rosaceae #0* Cotoneaster glaucophyllus (Cotoneaster) #0* Duchesnea indica (Wild Strawberry) * Fragaria ananassa (Strawberry) *Malus sp. (Apple) * Prunus sp. (Plum and Cherry) #0* Pyracantha angustifolia (Orange Firethorn) *Pyrus sp. (Pear) #0* Rhaphiolepis indica (Indian Hawthorn) *Rubus x loganobaccus (Loganberry) Rubiaceae Morinda jasminoides Rutaceae * Citrus sinensis (Orange) Solanaceae #0* Solanum pseudocapsicum (Madeira Winter Cherry) Sterculiaceae 0+ Brachychiton acerifolius (Iawarra Flame Tree) Verbenaceae #0* Lantana camara (Lantana) Vitaceae 0* Parthenocissus sp. (Virginia Creeper) * Vitis labrusca (Grape) * = introduced to Australia + = indigenous to Australia but not to the Thornleigh area 0 = present in the forest # = reproductive in the bushland No notation = native to the area RESULTS Pellets Occurrence The smallest number of pellets collected occurred in the autumn months (Table 2), a period which coincided with a seasonal decrease in the number of birds visiting the site (Buchanan 1983). Only eight pellets were collected in April, a deposition rate of 0.05 per square metre per month. The highest number occurred in September when 132 were collected, a deposition rate of 0.9 per square metre per month. The decrease in the number of pellets collected in July cannot be explained. The average width of coherent pellets was 15mm with a standard deviation of 0.32. The average length of coherent pellets was 27mm with a standard deviation of 0.64. The maximum number of non-coherent pellets (Fig. 3) was recovered over the late autumn and winter months of April, May, June, July and August, when the pellets lacked animal remains to bind the seeds together (Fig. 4). The maximum number of coherent pellets was recorded in October (58%) and November (71%) when pellets contained large amounts of mulberry (Morus alba). The majority of these multiple fruits were relatively intact. The data collected over 25 months showed that feeding is concentrated on one or a few items during the time needed to produce a pellet (Fig. 2). Almost 40% of pellets contained one component and more than 70% contained one or two. PROC. LINN. SOC. N.S.W., 111 (4), 1989 R. A. BUCHANAN 40 30. 20. pellets 10 3 4 NO components - 2 yrs_ data 5 Deal Fig. 2. Percentages of pellets containing different numbers of different components for the 25 month period. % pellets PARTLY COHERENT Fig. 3. Average monthly percentages of coherent, partly coherent and non-coherent pellets. PROC. LINN. SOC. N.S.W., 111 (4), 1989 248 CURRAWONGS AND WEED DISPERSAL 100. O O Q O 7 a Q O ® Oy H 80. 0 e O C) ‘a } Wi, rh es / Daa! LC] / D 60 ‘ % CL] Pellets 4 . O re) plant 20, @——® animal O—y introduced plants month Fig. 4. Average monthly percentages of pellets containing plant parts, introduced plants and animal parts for the 25 month period. Components and Seasons Plant Content The percentage of pellets containing plant parts remained high (79-98%) through- out the two years (Fig. 4). Fruit and seed was the major component (Table 3) but leaves and flowers were recorded in 3% and 0.6% of pellets examined. Almost half (22) of the 46 recorded fruits and seeds were present in less than 10% of the pellets in any one month (Table 3). Twenty-one species were abundantly represented, (Fig. 5). Oleaceae, including Ligustrum sinense, L. lucidum and Olea africana were present in over 20% of pellets from May to September and peaked at 74% in July. L. sznense was the most abundant and was present in over 20% of pellets for three months of the year (Fig. 6). The consumption of Olea africana was highest in the early winter months but never peaked above 20% and L. lucidum was the most important in the later part of winter (26% in August). Two other common weeds of urban native vegetation, Ochna atropurpurea and Cinnamomum camphora, were a prominent part of the diet in summer and autumn respec- tively. Insufficient sampling may have exaggerated the importance of Cinnamomum PROG. LINN. SOC. N.S.W., 111 (4), 1989 R.A. BUCHANAN 249 camphora in April of the second year — only eight pellets were collected. Morus alba dominated the diet in October and November and Pyracantha angustifolia was consumed for several months of the year (Fig. 5). 4st year and year % pellets 80 60 40 20 20 40 60 80 Oo Elaeocarpus reticulatus |___———] S_— Ligustrum lucidum* MM alban® het ode ee ESE IM alba J [ Persoonia pinifolia | ————__F Ochna atropurpurea’ a Ochna atropurpurea*{ si Ss CO. atropurrpureae Prunus sp..|_____—*|_ ~_| Solanum _pseudocapsicum’* Malus sp."|__——d| ds Phenix sp.” O. atropurpurea" |__| Vitis labrusca’ Lantana camara* F | Ficus carica’____ ———————_—sS's«@Ficus carica® Vitis labrusca® |_| SSCCSd’:séPhoolenix sp’ | Diospyros kaki’ =i‘ _—————_—*«|:«#Pyracantha angustifolia’ Pyracantha angustifolia"|__| S. pseudocapsicum* |_| M. alba’* P. angustifolia” Cinnamomum camphora® ee S. pseudocapsicum * M Melia azedarach’ Syzigium oleosum*|__ Cd Ss. angustifolia” J Ligustrum_sinense * Ligustrum_sinense* O._africana* |___——*d|s«~Pittosporum undulatum J Llucidum’ [Ys ucidum* L. lucidum * |_| L. sinense’ | L.sinense’ | CSC. cidum A |____J} P angustifolia® E. reticulatus |__| Elaeocarpus reticulatus L. sinense* |___|_| L. sinense’ |__| L. lucidum* |__| Smilax glyciphylla Fig. 5. The two plant species most frequently present in pellets for each month, showing changes between the first and second years of the 25 month period. Introduced plants were present in 50-90% of pellets in all but one month (Fig. 4). Twenty-one of the 36 introduced plants recorded are present in the bushland and at least 15 of these produce seed in this situation. Only 10 of the 46 species were native to the Thornleigh area. The most important of these were Elaeocarpus reticulatus, Persoonia pint- folia, Pittosporum undulatum and Smilax glyciphylla (Fig. 5). Only Elaeocarpus reticulatus was PROC. LINN. SOC. N.S.W., 111 (4), 1989 250 CURRAWONGS AND WEED DISPERSAL the dominant native species in any month (September). Prttosporum undulatum, which has a similar habit and similar habitat requirements to Ligustrum sinense and which is dispersed over the same months as the Oleaceae, was sparingly eaten. 80, w---@ Oleaceae w Y 60. oO—o Ligustrum sinense % e @ Olea africana pellets oO — o Ligustrum lucidum 40. 20. month Fig. 6. Average monthly percentages of pellets containing Oleaceae, Ligustrum sinense, Ligustrum lucidum and Olea africana. The data in Table 4 indicate that the fruit of the most favoured species have an average diameter greater than 5mm. Fruits of several species (Morus alba, Ficus sp., Solanum pseudocapsicum and Pittosporum undulatum) contained many seeds; but, the majority of fruit eaten contained only one or two seeds. For this latter group, the smaller the fruit is, the higher the number of seeds contained in a pellet. Fruits 5-6mm in diameter were represented by an average of 8-42 seeds per pellet while large fruit such as Persoonia pinifolia (14mm), Syzygium paniculatum (14mm) and Melia azedarach (13mm) were only represented by 2-5 seeds per pellet. Flesh and skin, but no seeds, of the very large fruited apple and persimmon were recorded in the pellets. The maximum number of seeds per pellet for Ligustrum sinense was 162, Lantana camara 131, Ligustrum lucidum 67, and Ochna atropurpurea 42. The major fruits in the currawong diet ranged in colour from bright orange (Diospyros kaki, Pyracantha angustifolia and Solanum pseudocapsicum) to green (Vits labrusca and Persoonia pinifolia) to black-blue for 7 of the 21 most frequent species. Fruit Selection Only ripe fruit was eaten or ingested by the pied currawongs. For green and pale- coloured fruit pied currawongs appeared to use means other than colour to determine ripeness. Just before Melia azedarach fruit first appeared in pellets, individual currawongs were observed to visit the tree; in each case the bird selected several fruits, held each one in the beak for a few seconds, and then dropped it. It is suggested that the birds were PROC. LINN. SOC. N.S.W., 111 (4), 1989 R. A. BUCHANAN 251 assessing the yellowish coloured fruit for ripeness by testing the softness of the pericarp with their beaks or tasting the flavour of exuded juices. TABLE 4 Fruit size and colour, number of seeds per fruit, average and maximum number of seeds in pellets Species Malus domestica Diospyros kaki fruit very large — only skin and flesh ingested, no seed may have many hundreds of seeds per pellet Phoenix sp. 1 3 Ficus sp. may have many hundreds of seeds per pellet Prunus nigra approx 1 1 20 Pittosporum orange only approx approx undulatum consume 50 200 seed Vitis labrusca Persoonia pinifolia Syzygium paniculatum Melia azedarach Solanum orange approx approx pseudocapsicum 60 300 Cinnamomum black 4 23 camphora Elaeocarpus reticulatus blue 22 Pyracantha orange approx angustifolia 60 Olea africana 8 Ligustrum sinense 6 36 Ligustrum lucidum 6 18 67 Lantana camara 5 131 Smilax glyciphylla 3) 54 PROC. LINN. SOC. N.S.W., 111 (4), 1989 252 CURRAWONGS AND WEED DISPERSAL Animal Content The percentage of pellets containing animal parts was highest (50-75%) in the warmer months but decreased to a low of 11-12% in the two coldest months of the year, June and July (Fig. 4). Remains of 7 major animal groups were recorded; these were Gastropoda, Arachnida, Diplopoda, Insecta, Amphibia, Aves and Mammalia. Vertebrate remains were rare in the collected pellets and only 14 contained bone fragments (Table 5). The fragments in seven of the pellets were too small for identifica- tion, but of the remaining seven, two were from birds, two from frogs, two from house mice (Mus musculus) and one was a chicken bone. The majority of the animals eaten were invertebrates — especially insects. Adult beetles, lepidopteran larvae and hymenopterans, — particularly bull-ants — were the most abundant remains (Table 5). The percentage of pellets containing bull-ants was high in the summer months, peaking at 47% in December, but fell to 0-4% during the winter months. This decrease was correlated with reduced bull-ant activity outside the nest (Fig. 7). TABLE 5 Animal remains identified in pellets Month and Number of Pellets Containing Item Categories of Animal Remains ), SES IMO VAS MiG]! AG SS = O= SNe Gastropoda (snails) 2 Z Arachnida (spiders) 1 Diplopoda (millipedes) 1 Insecta Ephemeroptcra (mayflies) 1 Blattodea (cockroaches) 1 1 Mantodea (praying mantids) 1 Dermaptcra (carwigs) 3 1 Orthoptera (grasshoppers & crickcts) 1 ar 1 1 2 1 Hemiptera (bugs) 2 Coleoptera (adult beetles) 30 16 9 8 10 3S & F 4 Ol BW sé Lepidoptera (moth & butterfly larvac) Daa! alle le ei ec sais) = 2 OR Soon |) other 1 7 saa \ ae ae 1 2, Hymenoptera (sawflies, wasps, bees & ants) Myrmecia sp. (bull-ant) 223 Oe ee On 3.20 52 26 40 other SOY el Se a TA ett aaa) 1 SIE 93 3 I @ Unidentified 0 12° 10 @ 8 22 lily te e20) (25s Bone Zar RMe E7 A alk ihre ae 2 2 1 Egg shell i 2 Number of pellets examined Y @7 42 47 GW) SY D7 BY GH 140 Br BH * = 3 months’ data DISCUSSION Pellets and Diet The examination of regurgitated pellets does not reflect the complete diet since only indigestible items are recorded (Dorst, 1971). The comparative volume of items in the pellets is also only a general guide to their importance in the bird’s diet. For example, remains of lepidopteran larvae were usually only represented by the head capsule even though the larvae may have contributed a significant portion of the day’s intake of food. On the other hand, favoured fruit items such as Elaeocarpus reticulatus, Olea africana, Ligus- trum sinense and L. lucidum have thin fleshy layers around a hard endocarp and lost very little of their volume after digestion. PROC. LINN. SOC. N.S.W., 111 (4), 1989 R. A. BUCHANAN DDS Despite the drawbacks outlined above the pellet studies reported here have aug- mented information on currawong diets in several ways. The importance of plant material in the omnivorous diet has been confirmed, but the wide diversity of fruits and seeds ingested — including many introduced species — has been documented in detail; the variety of animals eaten has also been demonstrated. Seasonal frequency in a number of animal and plant remains, and the small number of dietary components in individual pellets constitutes further evidence that currawongs are opportunistic feeders. Myrmecia sp. bull-ants pellets J month Fig. 7. Percentages of pellets containing bull-ants (Myrmecza spp.) in the 25 month period and the average number of bull-ants recorded at nest surfaces for one year. Fruit Selection and Dispersal The utilization of a large number of introduced plants as food resources by the currawong has wider implications, because of the potential for dispersal of weed species. Dispersal is only possible if the seed is undamaged by ingestion. Many seeds included in the pied currawong pellets were identified by germinating them and viability was high. Ligustrum sinense viability ranged from 83-91% (three samples of 50, 46 and 12 seeds), Pittosporum undulatum 100% (8 seeds) and Toxicodendron succadaneum 100% (6 seeds). The Oleaceae (Ligustrum sinense, L. lucidum and Olea africana) seem to be ideally suited to dispersal by pied currawongs. The prolific crop is produced in winter at a time when pied currawongs form large feeding flocks in the warmer parts of their range, including Sydney (Buchanan, 1983). The black-blue colour of the fruit is one of the colours preferred by birds (Turcek, 1963). The size of the fruit is at the lower limit of the size taken by pied currawongs so that large numbers of fruit, and hence seeds are con- sumed. The ratio of fruit to animals eaten also increases in the winter months so that a higher proportion of fruit may be eaten than in summer. Deposition of at least some of the seed in suitable conditions for germination and establishment (i.e. along creeks and PROG. LINN. SOC. N.S.W., 111 (4), 1989 254 CURRAWONGS AND WEED DISPERSAL rivers) 1s ensured by the fact that pied currawongs usually regurgitate pellets after drinking (Robertson, 1969; Clyne, 1980). The average number of seeds deposited per square metre can be large; in June 1977 approximately 12 Ligustrum sinense seeds were dropped on every square metre of the collection site. The density of deposition away from the collection site would be much less, but the common occurrence of clustered Ligustrum sinense, L. lucidum and Cinna- momum camphora seedlings in natural vegetation confirms that seed dispersal by birds is important. Some fruit appeared to be rejected by the birds. For example, Cotoneaster glaucophyllus is abundant in the study area and carried a prolific crop of fruit each autumn and winter, but was never present in more than 10% of pellets in any one month (Table 3). Rejection of Cotoneaster glaucophyllus cannot be explained. It was not on the basis of the thickness of the flesh (1.4mm) as this was greater than for Elacocarpus reticu- latus and the Oleaceae (0.8-1.2mm). Fruit size was within the range of the most frequently eaten species (Table 4) and by the colour it should have been selected (Turcek 1963). As well as the dispersal of obviously palatable items, pied currawongs may also distribute the seed of woody fruit. The fruit of Eucalyptus resinifera was found in one pellet. Even if the inclusion of such fruit and viable seeds 1s a rare event, the possibility for dispersal of minor dietary items is important. ACKNOWLEDGEMENTS My thanks are due to S. Buchanan for helping to collect the pellets and for the typing. The National Herbarium (Sydney) and the Australian Museum identified some of the more difficult items. My thanks are also due to H. Recher who commented on the manuscript. References AGNEW, N. V. 1., 1922. — Further notes from Peel Island, Moreton Bay, Queensland. Emu 21: 131-137. ANON., 1920. — Birds and insects. Emu 19: 248-251. ANON., 1927. — Birds and prickly pear. Emu 26: 203-206. BLAKERS, M., Davies, S. J. J. F. and REILLY, P. N., 1984. — The Atlas of Australian Birds. Melbourne: Royal Australasian Ornithologists Union, Melb. Univ. Press. BourkE, P. A.. 1949. — The breeding population of a thirty-five acre “Timber Paddock. Emu 49: 73-83. BRAVERY, J. A., 1970. — The birds of Atherton Shire, Queensland. Emu 70: 49-63. BUCHANAN, R. A., 1983. — Seasonal variations in population size of pied currawongs Strepera graculina at Thornleigh, Sydney. Aust. Birds 17: 49-55. CHENEY, G. M., 1915. — Birds of Wangaratta district, Victoria. Emu 14: 199-213. CLELAND, J. B., MAIDEN, J. H., FERGUSON, E. W. and Musson, C. T., 1918. — The food of Australian birds. N.S.W. Dept Ag. Sci. Bull. 15. CLYNE, D., 1980. — Wildlife in the suburbs. Up with privet. Sydney Morning Herald. 10 June: 7. COMMONWEALTH SCIENTIFIC AND INDUSTRIAL RESEARCH ORGANIZATION., 1970. — The Insects of Australia. Melbourne: Melb. Univ. Press. Cooper, C. D. and Cooper, R. M., 1981. — Observations on the food sources utilized by pied currawongs. Aust. Birds 15: 50-52. Currey, J. E. B., 1966. — Reflections of the Colony of New South Wales. Melbourne: Lansdowne Press. DorstT, J., 1971. — The Life of Birds. I. London: Weidenfeld and Nicolson. EDWARDS, H. V., 1922. — A south-coastal Selbourne. Emu 21: 42-48. KEAastT, A., 1958. — The influence of ecology on variation in the mistletoe-bird (Dicaewm hirundinaceum). Emu 58: 195-206. Lea, A. M. and Gray, J. T., 1936. — The food of Australian birds. An analysis of the stomach contents. V. Emu 35: 335-347. MARSHALL, A. J., 1935. — On the birds of the McPherson Ranges, Mt Warning and contiguous lowlands. Emu 35: 36-48. Morse, F. C., 1922. — Birds of the Moree district. Emu 22: 24-36. PROC. LINN. SOG. N.SW.,, 111 (4), 1989 R. A. BUCHANAN 255 READSHAW, D. L., 1965. — A theory of phasmatid outbreak release. Aust. J. Zool. 13: 457-90. READSHAW, J. L., 1968. — The distribution, abundance and seasonal movements of the pied currawong, Strepera graculina (Shaw), an important bird predator of Phasmatidae, in eastern Australia. Aust. J. Zool. 16: 37-47. RECHER, H. F., 1976. — Reproductive behaviour of a pair of pied currawongs. Emu 76: 224-226. Roserts, N. L., 1942. — The winter flocking of the pied currawong. Emu 42: 17-24. ROBERTSON, J. S., 1969. — Measurements, weights and notes on pied currawongs. Aust. Bird Bander 7: 7-10. Rose, A. B., 1973. — Food of some Australian birds. Emu 73: 177-183. SMITH, L., LARKINGS, D. and PEGLER, J., 1984. — Garden plants attractive to birds. Aust. Birds 18: 17-27. SPECHT, R. L., 1970. — Vegetation. Jn LEEPER, G. W. (ed.) The Australian Environment. Melbourne: C.S.I.R.O., Melb. Univ. Press. SPECHT, R. L., ROE, E. M. and BOUGHTON, V. H. (ed.), 1974. — Conservation of major plant communities in Australia and Papua New Guinea. Aust. J. Bot. Supplement No. 7. TURCEK, F. J., 1963. — Colour preference in fruit and seed-eating birds. Proc. XIII Intern. Ornithol. Congr: 285-292. VELLENGA, R. E., 1966. — Notes on the pied currawong. Aust. Bird Bander 4: 6. WALSH, J. E., 1965. — Notes on the pied currawong. Aust. Bird Bander 3: 53. WIMBUSH, D. J., 1969. — Studies on the pied currawong Strepera graculina in the Snowy Mountains. Emu 69: 72-80. PROC. LINN. SOC. N.S.W., 111 (4), 1989 eee cera ’ “APN ie ive SA eee tis : } Newb ce uot ty Wy val a “¢ 4 ee y a bre Mine eu. ‘vee i nite ‘a ty yy, pA yeaa MW aa hag te nin meh. Jeane hy Hh si a a oh a a we i a a we He WMG | ey Ale ; iy et ye Maat vey EL a Ny . 4it Vee ine anes m PN oe ak +) fi xv eh yee ah | i nen ATi Dr ae ~ Awana i ti“ ans 7 itaa Hi cn me NY CR eee bs th 4! Lun sar alone vey ed sie Me wae Po nh | i‘ wah kya Bs (iia ee eRe in AOR Lar ta B ‘ i TAY | ae Ra LE Als iii, i, 4 ‘ Bat N (hy! Ke Pe) na i nina f vi A te Pe neste y tno wine a an iui fea er ae ewe Di Me Vik ee ys , y SNP as ire ORE MRT ab Te Tha: BEV ALS Rete OR fig 7g a) Uy a ae o.. hes ws hie ra kal Gedy played lon cahan he pele bib pad dy tes Dts A Fda) cath i a Ta Vane ‘ial PA GATE: 8 hah ee LA me (un a albricails oe i 1 Yah whe i" ivy ie ma) i fk. a KS “Th i sth ute ; Sct i Mt ted a es Ache ‘ fr TaN TH iy WY eae PAR eb (An am UP i Ns Ty tT slg Woz i ri Ong, i? et AY af ( a ei tat) Nii ‘iio neice ved mit ee Ati ie s' Der ‘ we 4 r Shi . bi NA Ae ie olen: Dae of ua Wid ‘ Nine New Deep-water Species of Echinodermata from Norfolk Island and Wanganella Bank, northeastern [asman Sea, with a Checklist of the Echinoderm Fauna FRANCIS W. E. ROWE Rowe, F. W. E. Nine new, deep-water species of Echinodermata from Norfolk Island and Wanganella Bank, northeastern Tasman Sea, with a checklist of the echino- derm fauna. Proc. Linn. Soc. N.S.W. 111 (4), 1989: 257-291. Nine new species comprising 3 holothurians, 2 echinoids, 3 asteroids and 1 crinoid are described from the deep waters surrounding Norfolk Island and Wanganella Bank, northeastern Tasman Sea. A checklist of the echinoderm fauna shows at least 123 species are now known for that region, which includes the nine new taxa and seven taxa identified only to the level of genus. Taxonomic notes are included in the checklist for the taxa Bathyplotes punctatus (Sluiter); Holopneustes inflatus Lutken in Agassiz; Ophiothrix (Acanthophiothrix) lepidus de Loriol; Fromia polypora H. L. Clark; Coscinastertas muricata Verrill; Astrostole rodolphi (Perrier); Oxycomanthus plectrophorum (H. L. Clark) and Cenolia spanoschistum (H. L. Clark). Francis W. E. Rowe, Division of Invertebrate Zoology, Australian Museum, PO. Box A285, Sydney South, Australia 2000; manuscript received 7 December 1988, accepted for publication 19 July 1989. INTRODUCTION Norfolk Island (29°S, 168°E) and Wanganella Bank (c. 32°32’S, 167°32’E) lie to the northwest of the northwest tip of New Zealand. A number of recent, published accounts describe, or include records of echinoderms from those regions (Pawson, 1965b; McKnight, 1967, 1968a,b, 1975, 1977; H. E. S. Clark, 1970, 1982; Baker, 1979, 1980; Rowe, 1977, 1985; Edgecombe and Bennett, 1983; Rowe and Albertson, 1987). During the course of an investigation of the systematic composition and zoogeographic relationships of the echinoderm fauna of New South Wales (Australia), Lord Howe Island and Norfolk Island (Tasman Sea), the author has examined echinoderm material held in the collections of the Australian Museum (AM), Museum of Victoria (MV), National Museum of New Zealand (NMNZ) and the collections of the New Zealand Oceanographic Institute (NZOI). This latter collection includes, particularly, a great deal of material from Norfolk Island and Wanganella Bank along the Norfolk Ridge. Previous publications have recorded nearly 70 species from those regions. The present study has revealed not only nearly double that number (123 species) can be recorded, but that 9 of those are species which are new to science and seven taxa are identified as far as genus. The purpose of this paper is to provide descriptions of the new species and present a preliminary, updated checklist of all the taxa now recognized. The checklist includes a citation of the reference recording the taxon from the vicinity of Norfolk Island or Wanganella Bank; a general distribution similarly with an appropriate refer- ence and/or indication to ‘this work’ indicating the present author’s view; depth range; brief taxonomic note for each of 8 species where a comment is required. New records from Norfolk Island or Wanganella Bank are denoted by an asterisk (*). For the pur- poses of this report an area delimited by the latitudes 28°-33°S and longitudes 167° - 169°E is used to include records from waters surrounding both Norfolk Island (28° - 30°S, 167° -169°E) and Wanganella Bank (>30°-33°S, 167° -169°E). PROG. LINN. SOC. N.S.W,, 111 (4), 1989 258 NEW TASMAN SEA ECHINODERMS SYSTEMATIC ACCOUNT Class HOLOTHURIOIDEA Family Holothuridae Holothurta ( Vaneyothuria) uncia n. sp. Fig. 1A-D Diagnosis: A species in the subgenus Vaneyothuria which is distinctively coloured uni- formly cream with a chocolate brown ring around each tube-foot and with the rim of the disc of the 3-dimensional table spicules smooth. Material examined: Holotype, AM J21696, 29°24.8’S, 168°10’E, off Norfolk Island, 342-360 m (NZOI stn 191). Description: The contracted holotype is about 135mm long and about 57mm wide at the middle of the body. There are 17 tentacles. The tube feet are in three bands along the ventral ambulacra. The ventral-lateral bands are formed by a zigzag alignment of the pairs of tube feet. The mid-ventral band comprises two discrete rows of paired tube feet, the rows spaced about 6-8mm apart. Dorsally the tube feet are more or less scattered, without linear arrangement. The body wall is relatively thin. The calcareous ring is stout and not unusual (Fig. 1A). Tentacle ampullae are present. The gonad comprises a large bunch of branched and unbranched tubules on the left hand side of the dorsal mesentery. It lies about 20mm from the anterior end of the body, and the tubules are up to 60mm long. The respiratory trees are not unusual. The gut is full of sand. Spicules of the tentacles comprise curved spinous rods. The largest rods, from the tentacle stalk, measure up to about 480 wm long x about 50 wm wide. The smallest rods, from the tentacle branches, measure 50-150 wm x 5-7.5 ym (Fig. 1B). Spicules from the dorsal and ventral body walls comprise tables and buttons. The disc of the tables are squarish to irregularly rounded, smooth-rimmed, 90-105 wm in diameter and with a single ring of 8-10 holes (Fig. 1C). The spire comprises four pillars, joined by one or two cross-beams. The crown of the spire is small (c. 20-25 pm diameter), with a variable number of small spines. The spire is between 70-90 ym in height. The buttons are usually smooth, irregular in outline and usually have three pairs of holes (Fig. 1D). Rarely one or two low knobs may occur along the midline. They measure between 52 wm long x 37 wm wide to 127 wm long x 97 wm wide, with the majority measuring 90 wm x 52 pm. Tables in the dorsal tube feet range up to 112 pm disc diameter, with up to 16 small, peripheral holes, either in a single ring or with some holes offset into a partial, second ring. Buttons are present, but in addition curved and straight supporting rods occur. An irregular end-plate is also present, c. 180 pm diameter. Spicules in the ventral tube feet are often of similar type but the tables range in size from disc diameter 50-90 um and spires from 40-55 wm in height. The buttons are often elongate (165 wm long x 52 wm wide), with up to 6 pairs of holes. Supporting rods are similar to those in the dorsal appendages and the end-plates range up to 400 wm diameter. Colour: Uniformly cream, with a narrow basal ring around the ventral tube feet and a much broader (3-4mm diameter) ring of chococolate brown around the dorsal tube feet. Etymology: Named for Uncza, the snow-leopard. Remarks: This species (uncia) is clearly related to H. (V.) integra Koehler and Vaney, of which H. neozaelanica Mortensen is a synonym (Rowe, 1969; Cherbonnier and Feral, 1981) and which is distributed from the Bay of Bengal, off Port Hedland, northwest Australia, the Philippines and New Zealand. The spiny-disced tables of integra easily separate the two species. Also, the colour of wncza is distinctive among known deeper- water species of Holothuria. PROC. LINN. SOG. N.S.W., 111 (4), 1989 FW. E. ROWE 259 Fig. 1. A-D Holothuria (Vaneyothuria) uncia n. sp. (holotype AMJ21696), A = calcareous ring, interradial and radial plate, B = rods from tentacle, C = tables from body wall, D = buttons from body wall; E-H Mesothuria (Penichrothuria) norfolkensis n. sp. (holotype AMJ21697), E = calcareous ring, F = rods from tentacle, G = tables from body wall, H = ‘reduced’ table from tube-foot; I-M Neothyonidium parvipedum n. sp. (holotype, AMJ21698), I = lateral view of left side of holotype, J = calcareous ring, K = rods from tentacle, L = tables from body wall, M = end plate from tube foot. PROC. LINN. SOC. N.S.W., 111 (4), 1989 260 NEW TASMAN SEA ECHINODERMS Family Synallactidae Mesothuria (Penichrothuria) norfolkensis n. sp. Fig. 1E-H Diagnosis: A species in the subgenus Penichrothuria which is uniformly creamy-grey in colour; tube feet are scattered over the body but most prominent along the ventral- lateral ambulacra; reduced tables few; normal tables with 4 pillars which are crowned with ring of numerous small spines. Material examined: Holotype, AM J21697, 28°57.9’S, 167°45.5’E, off Norfolk Island, 392-423m (NZOI stn P35). Description: The holotype measures about 165mm in length and about 67.5mm in width. It has 17 tentacles. The tube feet are scattered over the body, but are most promi- nent along the ventral-lateral ambulacra. There are no tentacle ampullae. The polian vesicle is single. The madreporite lies to the right hand side of the dorsal mesentery. The gonad is a single tuft on the left hand side of the dorsal mesentery. It comprises some 25 tubules, each ending in a number of bifid lobes giving the gonad the appearance of a bunch of grapes. The water vascular ring 1s separated from the posterior end of the calcareous ring by about 15mm. The gut is packed with fine sand. The calcareous ring is stout, with radial plates deeply notched posteriorly (Fig. 1E). Spicules from the tentacles comprise more or less straight, or slightly curved, spiny rods. The largest, from the stem, range in length and width up to about 400 ym x 75 pm, respectively. The smallest rods occur in the branches of the tentacles and range between about 60-120 wm in length (Fig. 1F). Spicules from the body wall comprise tables only. Supporting rods and perforated disc plates are additionally present in the tube feet. Rarely in the tube feet are there small, reduced tables. The tables of the dorsal body wall have a disc diameter of 65-130 pm (Fig. 1G). There is either a single peripheral ring of 8-10 holes or an inner ring of 8 larger holes alternating with a partial or complete outer ring of 4-10 holes. The spire comprises four pillars with one cross beam. The spires are 52-75 wm high and crowned with a ring of numerous small spines. The tables of the ventral body wall are slightly smaller, with disc diameter up to about 112 um. The tables of the dorsal tube feet are smaller with disc diameters ranging from 50-90 um and spires up to 52 wm high. Other- wise, the tables have similar appearance to those of the body wall. The supporting rods are curved, perforated terminally and either side centrally. They are up to 187 um long. The end plates are up to 300 wm in diameter. Small, irregular, reduced tables are present, but very few in number (Fig. 1H). Spicules of the ventral and ventral-lateral tube feet are similar, in all respects, to those of the dorsal tube feet, except that the end plate of the ventral-lateral tube feet have a diameter up to 550 pm. Colour: Uniformly creamy-grey. Etymology: Named for the locality of discovery, Norfolk Island. Remarks: The form of the tables, with a complete ring of numerous small spines at the apex of the spire easily separates this species from all others, including M. (P) verrilli (Theel) and M. (P.) carnosa Fisher, in the subgenus Penichrothuria. Were it not for the occurrence of ‘reduced’ tables, albeit few, in the tube feet, I would consider M. (P.) norfolkensis closely related to M. (Allantis) intestinalis (Ascanius), though separable from it on the shape of the normal tables (the spire is more slender in intestinalis). Heding (1942) is quite emphatic that the absence of ‘reduced’ tables gives a clear limit to the subgenus Allantis in which he placed intestinalis. Because of the apparent rarity of the ‘reduced’ tables in M. (P.) norfolkensis, a fresh assessment of this character would be usefully under- taken. This is particularly relevant because, in their rarity, these tables can easily be PROC. LINN. SOC. N.S.W,, 111 (4), 1989 F. W. E. ROWE 261 overlooked. It is on this somewhat shaky character alone that norfolkensis is subgeneri- cally separated from its apparent nearest relative, intestinalis. Family Phyllophoridae Neothyonidium parvipedum n. sp. Fig. 1I-M Diagnosis: A species of Neothyonidium with a caudal process, small tube feet in double or irregular quadruple rows on the ventral ambulacra, more or less scattered dorsally; calcareous ring tubular, polyplacous, radials with very long posterior processes, inter- radials almost as long as the processes of the radials; two-pillared tables in body wall. Material examined: Holotype, AM J21698, 32°36.32°S, 167°30.7’E, Wanganella Bank, 126m (NZOI stn P4). Description: The contracted holotype is about 15.25mm long and about 6mm wide at mid body. The body is bluntly tapered anteriorly, but narrowing to a distinct caudal process (tail) posteriorly. The tail is dorsally directed, about 2.5mm from the ventral, backward projection of the body (Fig. 11). The tube feet are small, in double rows in the ambulacra anteriorly and posteriorly. Mid-ventrally the tube feet are widely spaced in somewhat irregular rows with 4 rows of tube feet in each ambulacrum. Dorsally the podia are fewer in number and without apparent regular arrangement. There are 20 tentacles, 5 pairs of larger alternating with 5 pairs of small tentacles. The calcareous ring is polyplacous, massive, tubular, about 13.25mm long. The radial plates are notched anteriorly, and have very long posterior processes. The interradial plates are pointed anteriorly and long posteriorly, extending to within about 3mm of the end of the ring. The interradials do not have posterior processes (Fig. 1J). The polian vesicle and madreporite are each single. The form of the gonad 1s difficult to determine but appears branched. It is preserved in a fused condition. Spicules of the tentacles comprise smooth, straight, curved, S-shaped or, rarely, X-shaped rods (Fig. 1K). The rods are terminally expanded and perforated. They range in size from 75 um long x 3.5 ym wide to 240 wm long x 17 wm wide. Spicules in the introvert comprise two-pillared tables. The disc is irregularly oval to squarish. It has 4 large central holes surrounded by up to about 26-30 smaller holes in one or two alternate rings. The disc reaches a maximum diameter of 112 um. The spire measures up to 75 wm in height, has a single cross-beam and each of the two terminally divergent pillars bears 2-3 spines at the tip. Spicules in the body wall comprise scattered, two-pillared tables (Fig. 1L). The disc of the tables range from 90 wm x 75 pm to 112 wm x 105 pm. The disc comprises 4 large, alternating with 4 small peripheral holes. Rarely, an outer, incomplete third row of 3-4 smaller holes is present. The spire comprises two pillars joined by a cross-beam. The tips of the pillars diverge and each is usually minutely bifid. The spires range from 40-60 ym in height. There are only end plates in the tube feet and these range in diameter from 75 pm (anteriorly and posteriorly) to 100 wm (mid-ventrally) (Fig. 1M). Colour: Uniformly greyish-white. Etymology: Named for its small tube feet. Remarks: This species is most closely related to N. hawazense (Fisher). It differs from hawauense in possessing a caudal process, in the slightly smaller disc plates from the tube feet, in possessing rods in the tentacles, and in the smaller number of perforations in the discs of the tables in the introvert. It is possible that parvipedum will be found to be conspecific with hawazense, but insufficient material does not permit such a conclusion at this time. I am not convinced by Heding and Panning’s (1954) synonymy of Fisher's PROG. LINN. SOG. N.S.W., 111 (4), 1989 262 NEW TASMAN SEA ECHINODERMS Fig. 2. Hapalosoma pulchrum n. sp. (holotype, AMJ21699), A = aboral view, B = oral view (h.d. = 67.4mm). PROC. LINN. SOC. N.S.W,, 111 (4), 1989 F. W. E. ROWE 263 (1907) second Hawaiian species N. alexandri with hawazense. There appears to be as close a relationship between alexandri and N. armatum Pawson, 1965c, from New Zealand as between hawazense and parvipedum. Class ECHINOIDEA Family Echinothuriidae Hapalosoma pulchrum n. sp. Figs. 2A-B, 3A-B Diagnosis: A species of Hapalosoma lacking aboral primary tubercles. Material examined: Holotype, AM J21699, 29°54.90’S, 44°.80’E, off Norfolk Island, 130-301m (NZOI stn P26). Description: The test is flattened, more or less circular in outline, with h.d. = 67.4mm (Fig. 2A-B). Apical system: Measures 13mm in diameter. Ocular plates are insert, anvil-shaped, contiguous with the more or less kite-shaped genital plates. The genital pores are large, indicating the probability of a female. A number (8-12) of small tubercles occur on a discrete convexity of the plates adapically to the genital or ocular pores respectively. The madreporite is large and prominent, extending over most of the plate. Apical plates each bear 1 or 2 tubercles (Fig. 3B). Ambulacra: There are 55 plates in each ambulacral column. Two demi-plates, each pierced by a pore pair, are associated with each ambulacral plate. The pore pair piercing the ambulacral plate lies adjacent to the interambulacral plate. The pore pairs form three columns. A large primary tubercle occurs on each second and/or third plate, internal to the pore pairs on the oral surface as far as the ambitus of the test. A smaller tubercle occurs on each plate between the outer demi-plate and pore-pair piercing the ambulacral plate. A second even smaller tubercle occurs regularly under the pore-pair piercing the ambulacral plate. The latter two tubercles form a regular double series to the ambitus in the oral side. Three or four additional smaller tubercles occur in a median transverse line across the plate. Above the ambitus there are no primary tubercles but a row of 6, diminishing to 0-1 adapically, small tubercles forming a median transverse row on the plates. Skin areas are very narrow, almost obliterated between the plates. The ambulacral width at the ambitus is 14.2-15.6mm (Fig. 3A). Interambulacra: There are 37 plates in each interambulacral column. The width of the interambulacrum is about 27mm at the ambitus. A large primary tubercle occurs on each of the first 14 plates from the edge of the peristome to the ambitus on the oral side. These tubercles form a regular series adjacent to the ambulacra. One or two additional tubercles, one usually close to the mid interambulacral margin of the plate, also occur, but irregularly on these plates. Up to about twelve minute tubercles occur scattered between the large tubercles. Above the ambitus, from about the 15th plate from the peristome, there are no large primary tubercles. Instead, a median transverse row of small, but equisized tubercles occur on each plate, the number diminishing from 13-14 at the ambitus to 0-1 adjacent to the apical system. Areas of skin between the plates are minimal (Fig. 3A). All tubercles are perforate and non-crenulate. Peristome: The peristome measures approximately 14mm in diameter. The plates carry flattened but club-shaped spines and pedicellariae. Pedicellariae: These are typical of the genus though there is a tendency for the shaft of the bifid-tipped, reduced dactylus pedicellariae to be perforated by up to three small holes. Colour: Test is pale green, the green being more intense on the ambulacral plates. The PROC. LINN. SOG. N.S.W., 111 (4), 1989 264 NEW TASMAN SEA ECHINODERMS 5mm Fig. 3. Hapalosoma pulchrum n. sp. (holotype, AMJ21699), A = plates ofambulacrum and interambulacrum, B = apical system. apical system is purplish. A wide purplish longitudinal band occurs in each ambulacral and interambulacral area orally, extending just above the ambitus as a purple patch. PROC. LINN. SOG. N.S.W., 111 (4), 1989 F. W. E. ROWE 2605 The larger oral spines are very pale greenish with 5-6 narrow purplish bands. Remains of the white ‘hoofs’ indicate they are not prominent. The smallest secondary spines are uniformly whitish to pale green as are the abactinal spines. Etymology: pulcher (Lat.) = beautiful, referring to its striking colour pattern. Remarks: This species differs from its only congeners H. pellucidum (A. Agassiz) and H. gemmiferum Mortensen primarily by the absence of aboral primary tubercles and spines. The pierced blade of the dactylus pedicellariae and the colour pattern may also be useful differences separating the species. The genus Hapalosoma was last reviewed by Mortensen (1935). Family Pedinidae Caenopedina alanbakeri n. sp. Figs. 4A-B, 5A-D Diagnosis: A species of Caenopedina with relatively short primary spines (1.37 x h.d.); apical system 33% h.d.; peristome 25% h.d.; interambulacral plates with large primary tubercle and prominent secondary tubercle the remaining plate surface covered with small tubercles of 2-3 sizes; milled ring of spines 10% wider than the spine; test pale pink, primary and secondary spines uniformly deep pink basally, lighter towards the tip. Material examined: Holotype AM J21700, 29°24.80’S, 168°13.20’E to 20°23.70’S, 168°13.80’E, off Norfolk Island, 570-578m (NZOI stn 192). Description: The test is circular at the ambitus, flattened aborally and orally, the sides strongly arched (Fig. 4A-B). The h.d. = 41 mm, v.d. = 26 mm. There are 19 coronal plates. Apical system: This measures 13.5mm in diameter (33% h.d.) (Fig. 5A). It is dicyclic with oculars all widely exsert. The male genital pores are small, horizontal, slit-like, with a channel extending from the pore to the lower border of the 8th interambulacral plate. The genital plate bearing the madreporite is not enlarged, the madreporite occupying a triangular, central portion of the septagonal plate. There are, more or less, two distinct groups of tubercles on each genital plate. A triangle of 10-11 tubercles occupies the periproctal edge of the plate while a further 9-10 tubercles form a more or less double transverse band across the plate, the remainder of the plate is bare (Fig. 5A). The oculars bear 5-6 small tubercles of which 4-5 form a transverse line straddling the ocular pore, the remaining tubercle occurring towards the inner edge of the plate. The periproct is covered by small plates, the anal aperture more or less centrally placed. Ambulacra: The ambulacra measure 6.2mm in width at the ambitus (= 35.8% of inter- ambulacra). Ambulacral plates are trigeminate, the pore pairs being in arcs of three on the oral surface from the peristome to the ambitus. Aborally, from about the ambitus to the apical system the pore arcs become more vertically aligned so that the pore pairs form a somewhat sinuous line. The middle component of each plate bears a conspicu- ous, perforate, non-crenulate, primary tubercle whose areole extends onto the plate components above and below it. These tubercles form a regular vertical series in each column, decreasing in size towards the apical system. Small secondary tubercles, of two sizes, occupy the remaining surface of the ambulacral plates (Fig. 5B). Interambulacra: The interambulacra measure 17.3mm in width at the ambitus. The plates each bear a large primary tubercle more or less in the middle of each plate. The tubercles form a vertical series. The areoles are large and confluent between plates on the oral surface. On the aboral surface, above the ambitus the areoles are separated adapically by a simple row of small, secondary tubercles. A prominent, secondary tubercle occurs near the mid-interradial edge of each plate, forming a second vertical series below the ambitus but a more or less zigzag series above the ambitus. The remain- PROC. LINN. SOC. N.S.W., 111 (4), 1989 NEW TASMAN SEA ECHINODERMS LOE Fig. 4. Caenopedina alanbakeri n. sp. (holotype, AMJ21700 oblique aboral view, B = lateral view (h.d. = 41mm). PROG. LINN. SOG. N.S.W., 111 (4), 1989 F. W. E. ROWE 267 ing surface of each plate is crowded with smaller, secondary tubercles of 2-3 different sizes. All tubercles are perforate and non-crenulate (Fig. 5B). C imm 5mm D 94um 60um Smm Fig. 5. Caenopedina alanbakeri n. sp. (holotype, AMJ21700), A = apical system, B = plates of ambulacrum and interambulacrum, C = milled ring of primary spine, D = spicules from tube-foot. (gg = genital groove). Peristome: The peristome measures 10.3mm (+ 25% h.d.). It is finely plated as well as containing the conspicuous buccal plates which bear pedicellariae. Spines: The longest spines are 56.4mm, slender, tapering, longitudinally striated with minute thorns. The milled ring is conspicuous but not widely produced so that it is only 10% wider than the spine (Fig. 5C). Secondary spines are similar to the primaries but much smaller, about 1/5-1/4 the length of the primaries. PROC. LINN. SOC. N.S.W., 111 (4), 1989 268 NEW TASMAN SEA ECHINODERMS Spicules: The tube feet contain spicules which are slightly curved, irregularly elongate, perforated plates or smooth rods with marginal perforations (Fig. 5D). Pedicellariae: These do not appear unusual for the genus, or to be distinctive, though the ophicephalous pedicellariae are rare on the test of the holotype. Colour: The dried test is uniformly a very pale pink (when cleaned with domestic bleach the test is white except for a persistent pink on the plates of the apical system and immediately adjacent coronal plates). The spines are not banded but are deeper pink basally, becoming paler along their length, the distal 4% to % of the spine being a pale yellowish/lime colour. The secondary spines are similarly coloured only even paler. The poison sacks of the globiferous pedicellariae are violet/purple. The tube feet are pale brown with the tip sienna brown. Etymology: This distinctive species is named for Dr Alan N. Baker, National Museum of New Zealand, who has contributed significantly to ophiuroid and echinoid taxonomy in this geographical region. Remarks: The genus Caenopedina A. Agassiz has been reviewed by Mortensen (1940). More recently two species have been described from New Zealand waters; C. novaezealan- diae Pawson (1964) and C. otagoensis McKnight (1968c). C. alanbakeri is immediately distinguished from its geographically nearest congener C. novaezealandiae Pawson by the colour; the size of the milled ring of the primary spines; the relative sizes of the apical system and peristome to h.d.; the relative number of coronal plates and tuberculation of the plates. Although the primary spines are not banded, C. otagoensis McKnight is other- wise distinguished from C. alanbakeri on similar comparative features to those of C. novaezealandiae. Despite the fact that the primary spines are only 1.37 x h.d., their slender form and the test plate ornamentation immediately distinguish C. alanbaker: from either of the short-spined C. pulchella (A. Agassiz and H. L. Clark) or C. superba H. L. Clark. These features clearly also distinguish C. alanbakeri from all other congeners. Class ASTEROIDEA Family Astropectinidae Tethyaster tangaroae n. sp. Figs. 6A-B, 7A-B Diagnosis: A species of Téthyaster which has stout tabulae (about twice as high as wide); actinal plates with 1-3 prominently elongate central spinelets; adambulacral plates and some actinal and oral plates usually bear a large, elongate bivalved pedicellaria. Material examined: Holotype, AM J21701, 28°57.90’S, 167°45.50°E, off Norfolk Island, 392-423m (NZOI stn P35), 2 paratypes, NZOI, 28°54.60°S, 167°44.20’E, off Norfolk Island, 390-402m, (NZOI stn P27). Description: The holotype measures R = 46.5mm,r = 11.3mm, br (at 2nd inferomar- ginals) = 10.8mm; R/r = 4.1, R/br = 4.3 (Fig. 6A-B). The arms are slender, tapering to a narrow tip which is occupied by a prominent, convex, longitudinally elongate terminal plate. The terminal plate has a bumpy surface when cleaned of the minute spinelets which cover it. The spinelets give the plate a shaggy, felt-like appearance. The abactinal plates have a six-lobed base and are tabulate. They reduce in height towards the tip of the arm. The tabulae are about twice as high as they are wide, somewhat waisted and of two sizes, mixed, which are arranged, more or less, in longiseries on the arms. There are thirteen series at the base of the arm, reducing to four series at the arm tip. It is possible to detect a carinal row of spaced, larger tabulae when the tabulae are cleaned of their spinelets. The centre of the tabulae is occupied by up to 25 short, blunt- tipped spinelets of mixed sizes, and the periphery by up to 25 slender spinelets. The madreporite is at about %r from the disc centre. It bears small spinelets on its PROC. LINN. SOC. N.S.W., 111 (4), 1989 F. W. E. ROWE Fig. 6. Tethyaster tangaroae n. sp. (holotype, AMJ21701), A = aboral view, B = oral view(R = 46.5mm). PROG. LINN. SOG. N.S.W., 111 (4), 1989 NEW TASMAN SEA ECHINODERMS V), SS, 4 AYSE ly o Sot Loe DY isd 4 ah! VW) AN U 1, Uj NN Me AY BA, ss ae y Z¢ Yj 2mm > N 2 2 7 US Y Zz INK hos W Mu (NN l <— mouth Imm Fig. 7. Tethyaster tangaroae n. sp. (holotype, AMJ21701), A= = superomarginal (sm) and inferomarginal (im) plates in interradial arc, B = 4th-6th adambulacral plates (ad) and adjacent actinal plates (ap), showing spines and pedicellariae (p). corrugated surface and is almost hidden by the spinelets of adjacent tabulae. The disc is more or less flat, though the radial regions, at the base of the arms, are slightly convex. PROC. LINN. SOC. N.S.W., 111 (4), 1989 F. W. E. ROWE 271 The papulae are single and occur between the tabulae over the whole abactinal surface, extending to the arm tips. The supero- and inferomarginal plates are similar in size and shape. There are 34 superomarginals and 35 inferomarginals, the distalmost 2-3 inferomarginals being very small and extending under the terminal plate each side. The marginal plates are verti- cally elongate, with a raised, flat-topped, rectangular median ridge which leave deep, straight-sided fasciolar channels between successive ridges. The flat-topped surface of the ridges are covered with low bumps, each bump bearing a short, stout spinelet. On the superomarginal plates these spinelets become elongate towards the edge of the ridge where together with those on the adjacent plates the spinelets span and cover the fas- ciolar channels between. The inferomarginal plates bear a vertical series of 3, proximally, reducing to 1 distally, large flattened, acicular spines across the ridged surface. Additionally, smaller slender, but otherwise similar spinelets cover the remain- ing surface with fasciolar spinelets, similar to those on the superomarginal plates, at the periphery of the ridge (Fig. 7A). The actinal surface is relatively small, with 4 rows of plates on each side and a supplementary row of three plates extending along the mid-interradial line between the oral plates and first inferomarginal plates (Fig. 6B). The first actinal row extends to about the 10th inferomarginal plate; the second row extends to about the 6th infero- marginal plate; the third row comprises 3 plates and the 4th is a single plate. The actinal plates have a central convexity which bears elongate spinelets, 1-3 central ones being more prominent, elongate, tapering spines. Several plates bear a large, elongate, bivalved pedicellaria, replacing one or more of the central spines. The adambulacral plates are somewhat wedge-shaped from actinal aspect, thereby projecting, slightly, into the furrow. There are 3-5 furrow spines, the middle spine being the longer and it is compressed. The actinal surface of the plates bear 3-4 central, elongate spines, one or two of which are replaced by a single, large, bivalved pedicellaria on many plates. The first two adambulacral plates each bear 2-3 such pedicellariae. The periphery of the plates bear a number of smaller, slender spinelets (Fig. 7B). The oral plates bear 6-7 furrow spines. The two adjacent apical spines are so closely appressed as to appear to be fused together. Along the median crest between the adja- cent oral plates is a row of 10-11 elongate spines, the 3rd and 4th of which are replaced by a large bivalved pedicellaria on several of the plates. Neither of the paratypes possess pedicellariae, otherwise they are similar in almost all other respects to the holotype. One paratype measures R = 34.6mm,r = 9.2mm, br = 7.6mm; R/r = 3.76, R/br = 4.6. It has 27 superomarginal and 30 inferomarginal plates. There are 3 actinal rows and 2-3 plates form the median interradial row. The second paratype measures R = 10.3mm,r = 4.2mm, br = 4.7mm; R/r = 2.45, R/br = 2.2. It has 14 superomarginal and 16 inferomarginal plates. The inferomarginal spines are hardly prominent. There are two actinal rows and 2 plates form the median interradial row. Remarks: T° tangaroae differs from its geographically nearest neighbour T’ aulophora (Fisher) principally by the stouter tabulae, form of the large bivalved pedicellariae, and spinulation of the actinal plates. 7’ tangaroae differs from the South African T’ pacez (Mortensen) by the actinal spinulation and pedicellariae. The discovery of a species of Téthyaster in the Tasman Sea is not an unexpected extension in range of this widespread genus which has been revised by A. M. and A. H. Clark (1954). PROC. LINN. SOC. N.S.W., 111 (4), 1989 DD NEW TASMAN SEA ECHINODERMS Fig. 8. Glyphodiscus mcknighti n. sp. (holotype, AMJ21702), A = aboral view, B = oral view(R = 23mm). PROC. LINN. SOC. N.S.W., 111 (4), 1989 F. W. E. ROWE DIB Family Goniasteridae Glyphodiscus mcknighti Figs. 8A-B, 9A-B Diagnosis: A species of Glyphodiscus with smooth, flat abactinal plates, smooth marginal plates and few papulae which are restricted to the radii on the disc. Material examined: Holotype, AM J21702, 28°42.30’S, 167°56.70’E, Norfolk Island, 475-450m (NZOI stn P46). Description: The holotype measures R = 23mm,r = 11.5 mm, br = 4.2mm (at 2nd inferomarginals); R/r = 2.0, R/br = 5.5. The abactinal area is pentagonal, slightly produced at the angles and sunken below the level of the superomarginal plates. The abactinal plates are smooth, flat, without crystal bodies. The plates are rounded-polygonal, the interradial plates being slightly larger than the radial plates. A row of narrow, transversely rectangular plates occurs adjacent to the superomarginals. A small, triangular madreporite occurs 4%r from the centre of the disc (Fig. 8A). The supero- and inferomarginal plates are similar in shape, size and number. They are block-like, smooth, longer than wide with a rounded dorsal-lateral or actinal-lateral edge respectively. There are 6 of each on each side of the disc and arm, the first 2 supero- and inferomarginals of adjacent radii respectively delimiting the pentagonal abactinal and actinal disc surfaces. The remaining 4 supero- and inferomarginals on each side of the arms unite across the arm along the median line. The terminal plate is small, with a convexity at its tip on either side of the actinal channel which houses the terminal tube foot. This channel is guarded by 3-4 minute granules. The actinal plates are flat, smooth, rather transversely diamond-shaped. They form a regular pavement arrangement (Fig. 8B). The abactinal, marginal and actinal plates are each surrounded by a single row of minute granules so that a double row of granules occurs between adjacent plates. Papulae are restricted in each radius abactinally and are delimited as 5 rounded, convex areas each comprising about 10 plates. Usually 1, occasionally 2, small spatulate pedicellariae occur at the edge of a number of actinal plates, also at the actinal edge of several of the inferomarginal plates (Fig. 9A). Pedicellariae are not present on the superomarginal or abactinal plates of this specimen. The adambulacral plates bear five laterally compressed furrow spines (Fig. 9B), behind which stands a row of 3 enlarged (subambulacral) granules. The remaining actinal surface of the plate bears 2-3 rows each of 3-4 smaller wedge-shaped granules which merge in size with those surrounding the actinal plates (Fig. 9A). The oral plates bear 7-8 furrow spines, 4-5 enlarged (suboral) granules and a tri- angular group of 8-10 wedge-shaped granules on the remaining actinal surface of the plate. Etymology: Named for Mr D. McKnight of NZOI who has described much of the Tasman echinoderm fauna. Remarks: I have placed this new species in the herein elevated subgenus Glyphodiscus Fisher on the grounds that the species characters are consistent with those outlined for the genus by Fisher (1917, 1919). I believe the smooth plates which lack glassy bumps (crystalline bodies), the complete ring of peripheral granules around all plates and the form of the adambulacral armature are quite reasonably sufficient to distinguish members of this genus from those in either Jconaster Sladen, Lithosoma Fisher or Astro- ceramus Fisher. G. mcknighti is clearly distinguished from its only congener G. perzerctus Fisher which PROG. LINN. SOC. N.S.W., 111 (4), 1989 274 NEW TASMAN SEA ECHINODERMS has roughened superomarginal plates, tumid peripheral abactinal disc plates and papulae distributed over the disc. In respect of size and shape, the two species are very closely similar. A 2mm 1mm — mouth 4th ad Fig. 9. Glyphodiscus mcknighti n. sp. (holotype, AMJ21702), A = spine and granule arrangement on adambulacral (ad) and adjacent actinal (ap) plates with pedicellariae, B = furrow spines (fp). Family Brisingidae Novodinia helenae Figs. 10A-B, 11A-C Diagnosis: A species of Novodinia with 12 arms, disc with papulae in groups of up to 10, PROC. LINN. SOC. N.S.W., 111 (4), 1989 F. W. E. ROWE 275 single madreporite, membranous, unskeletonised genital region of arm, and single adambulacral spine. Material examined: AM J21703, 29°20.20’S, 168°10.79’E, off Norfolk Island, 308m (NZOI, stn 194). Description: The holotype has 12 rays; dd = 14mm, R = 75mm, br = 3.8mm (at base), 5.5-5.9mm (at widest part of genital expansions 12mm from base of ray), 3.0mm (at %R); R/r = 10.7 (Fig. 10A-B). The disc is circular, 3.7mm high, flattened abactinally. It is covered by overlapping scale-like, and convex abactinal plates which form an open reticulum in which groups of up to 10 (occasionally single) papulae occur. Sharply pointed spines (up to 1.2mm), wreathed almost to their tips with crossed pedicellariae (Fig. 11C) occur on a number of the convex plates. These spines may stand singly, in pairs or triplets. In the latter two cases the spines are united by a web across which the pedicellariae span. The genital region, at the base of the arm, is covered by a thin, unskeletonised membrane. This region is crossed by 4-5 complete costae. The costae comprise the marginal plate each side of the arm, linked more or less regularly by 7-10 rod-shaped abactinal plates. Costae 1 and 2 may be irregularly linked by a few abactinal plates extending mid-dorsally between them. Most of the plates of the costae, except for the marginal plates of the first two costae, bear a single, sharply pointed, slender spine. Most of these spines are encased in a gland-like sheath of pedicellariae. Beyond the 4th- 5th costae the spine bearing marginal plates and abactinal plates form incomplete costae, that is to say a marginal and 4-5 small abactinal plates project dorsalwards along the side of the arm on each side but do not link over the abactinal surface. The link is made instead by a wide band of crossed pedicellariae. The abactinal surface of the arm is then covered by a very thin membrane for the rest of the length of the arm. Abactin- ally, between each of the incomplete costae a rounded patch of pedicellariae also occurs along the arms. The incomplete costae occur opposite every 4th-5th adambulacral plate along the arm (Fig. 11A). The adambulacral plates are block-like, wider than long and each bears a large, cylindrical spine (2.5mm) which at least for the length of the genital area are flared into a bi-quadrifid tip. Beyond the genital area these spines are slender and pointed. These spines bear numerous pedicellariae, but these are confined to the outer surface and do not ensheath the spines. The first adambulacral plate of each arm is fused across the interradial line to its adjacent neighbour. However the first two adambulacral plates on each side of each arm are united by a syzygy. Thereafter the plates are united by muscle blocks, the interstices between each successive pair of adambulacral plates being half the length of the plates themselves. The actinosome is 8mm in diameter and the mouth is 3.6mm wide. The peristome is thin but translucent. The oral plates are as usual for the genus, with lateral processes meeting mid-radially. In well-developed oral angles each plate bears 5 spines in a fan- shaped, marginal arrangement. The innermost spine is usually very small, rarely is it long and bearing pedicellariae. The second spine is always long and pointed and bears pedicellariae. The remaining 3 spines are small, non-pedicellariae bearing and decrease in size towards the furrow. Where arms are being regenerated, the mouth angle plates bear fewer spines (3-4) of which the elongate pedicellariae bearing spine is in the apical position. A small additional spine can be seen to develop in the interradial side of this spine in angles with more advanced regeneration. There are no spines on the actinal surface of the oral plates. The furrow margin of each oral plate is excavate to accommo- date the first pair of tube feet (Fig. 11B). The madreporite (1.2mm diameter) is small, subtuberculate, coarsely furrowed. PROC. LINN. SOC. N.S.W., 111 (4), 1989 276 NEW TASMAN SEA ECHINODERMS Fig. 10. Novodinia helenae n. sp. (holotype, AMJ 21703), A = aboral view, B = oral view (dd = 14mm, R = 75 mm). PROC. LINN. SOG. N.S.W., 111 (4), 1989 F. W. E. ROWE BT 7mm 120pm ad ads Fig. 11. Novodinia helenae n. sp. (holotype, AMJ21703), A = costae (c) in genital region of arm, B = oral plates (op) and arm bases with spine arrangements, C = value of crossed pedicellaria (ad = adambulacral plate; ads = adambulacral spine; tf = tube-feet). Etymology: Named for Dr Helen E. S. Rotman (nee Clark) who has contributed to our knowledge of Tasman and Antarctic asteroids. Remarks: The new species clearly belongs in the genus Novodinia Dartnall et al., as most recently redefined by Downey (1986). Only two species of Novodinia are recorded from the Tasman/New Zealand region. N. australis (H. L. Clark), from southeastern Aus- tralian waters and N. novaezealandiae (H. E. S. Clark), from off the Chatham Islands, east of New Zealand. N. helenae is immediately distinguished from each of these species by a number of characters: firstly, low arm number (12) as opposed to 14-16 (australis) or 18 (novaezealandiae); secondly, abactinal skeletal arrangement of the disc with groups of papulae (in each of australis and novaezealandiae the skeletal network is close and papulae occur singly); thirdly, the genital area of the arm is membraneous in N. helenae but plated in the other two species. The number of madreporites (1 in Aelenae but 4 in australis) and the number of adambulacral spines (1 in Aelenae but 3-4 spines in novaezealandiae) are also features of distinction. Arm number and ornamentation of disc and genital regions readily distinguish N. helenae from its other congeners. PROC. LINN. SOC. N.S.W., 111 (4), 1989 278 NEW TASMAN SEA ECHINODERMS Class CRINOIDEA Family Antedonidae Nanometra duala n. sp. Fig. 12A-E Nanometra johnstont. McKnight, 1977: 136 (non N. johnston: John) Diagnosis: A species of Nanometra with a pair of wing-like extensions to the segments of P, and subsequent pinnules. Material examined: Holotype, AM J21704 and 2 paratypes, NZOI, 29°20.20’S, 168°10.79’E, off Norfolk Island, 308m (NZOI, stn 194). Description: The holotype has 10 arms broken at about 20br and measuring about 10mm in length. The estimated length of the arms is not more than 20mm (Fig. 12A). The cirri are XXIV-X XVI, 19-22. The first 2 segments are twice as broad as they are long, the third segment is 1.5 x as long as it is broad, 4th-5th segments twice as long as broad, 6th segment 1.75 x as long as broad, 7th segment 1.25 x as long as broad. Distally, the segments are as long as they are broad. The segments do not bear a dorsal spine but the distal end is expanded (Fig. 12A-B). The centrodorsal is conical, with a rugose apex. The cirri are arranged in vertical rows of 2 or 3 (Fig. 12B). The radials are narrow, almost hidden, with a tubercle at each of the exposed corners of the plate. The 1Br, is twice as broad as it is long, with the proximal and distal edges more or less straight but with the distal edge everted. There is a rounded or chisel-shaped tubercle arising on each side of the ossicle and one or two spinulose tubercles below each of these on the dorsal-lateral surface of the ossicle. The axillary (1Br,) is triangular, slightly broader than long (1.25:1). The proximal border is slightly convex, the distal border everted and spinulose. There are one or two spinulose tubercles occurring on the lateral surfaces of the ossicle (Fig. 12B-C). Br, is quadrate, about 2-2.5 times as wide as long. Up to 4 spinulose tubercles may occur on the lateral surface of the ossicles. Remaining brachials are more or less elongate, wedge-shaped, with the distal edge everted and spinulose (Fig. 12C-D). Syzygies occur at 3+4 and usually 9+10, but occasionally the second syzygy occurs at 8+9 or 11+12 or 13+14. The third syzygy occurs at 14+15 or 15+16. P, is the longest pinnule and the stoutest. It is 3mm long, comprising 11 segments. The first segment is broader than long, the second is quadrate, the remaining segments are up to 2 times as long as broad. P,, is about 1.5mm long and comprises 6 segments. P, is smaller and more slender than P,, is about 2mm long and comprises 9 segments. The segments of these pinnules are strongly everted and spinulose on their distal edge. P, is the first gonadal pinnule. On P, and subsequent pinnules (for the length of the broken arms) the lateral edges of the 2nd to 4-5th segments are expanded into thin, wing-like processes, giving the pinnules a very characteristic form (Fig. 12D). One paratype is very similar indeed to the holotype in being a relatively intact, if broken armed, specimen. The second paratype comprises the calyx of a specimen with many broken arm pieces. The form of the segments of the pinnules is, however, unmistakable in uniting the specimens within a single species. Etymology: duo = two; ala = wing (Lat.) referring to the 2-winged appearance of the pinnular segments. Remarks: The very distinctive form of the pinnule segments immediately distinguishes this species from its congeners N. johnstoni John (from S.E. Australia); N. clymene A. H. Clark (from the East Indian region) and N. bowers: (A. H. Clark) (from southwestern Japan), which have been reviewed by A. H. Clark and A. M. Clark (1967). McKnight’s PROC. LINN. SOC. N.S.W., 111 (4), 1989 F. W. E. ROWE 279 (1977) record of N. johnston: from the same locality (NZOI stn 194) refers instead, I believe, to the new species N. duala. Smm 2.5mm 2mm Fig. 12. Nanometra duala n. sp. (holotype, AMJ21704), A = lateral representation of holotype, B = cirri, centrodorsal and 1Br, C = 1Br and arm bases, D = P3 with cross-section of pinnule segment showing wing- like extensions. ACKNOWLEDGEMENTS The author wishes to express his gratitude to the following people for allowing him unlimited access to their collections and for the loan of material: Messrs D. McKnight and W. L. Main (New Zealand Oceanographic Institute, Wellington); Drs A. N. Baker and H. E. S. Rotman (National Museum of New Zealand, Wellington); Sue Boyd (Museum of Victoria, Australia). Ms P. Berents and Ms Lynne Albertson (Australian Museum) are thanked for preparing figures and Ms Berents for reading and comment- ing on the manuscript. Mr. A. Farr (Australian Museum) for photographing the specimens. Ms Jan Howarth is thanked for typing the manuscript. This project has been PROC. LINN. SOC. 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A., 1982. — Sea-stars (Class Asteroidea). Jn SHEPHERD, S. A. and THOMAS, I. M. (eds.) Marine Invertebrates of Southern Australia Part 1. Adelaide: South Australia Government, pp. 400-418, figs 10.3-10.9, pl 30, fig. 2. PROC. LINN. SOC. N.S.W., 111 (4), 1989 NEW TASMAN SEA ECHINODERMS 282 ‘(GZ6]) uosue yy ‘ueijodousor) = t (J904,],) va9vj020 auosoujav7 “Ty aepiuosounoey Apimey FI9-OIFE ‘sajojddyjog yo sutAuouAs se ‘utotoy ‘Ou Aq popsedos ose sayrsdqiaydyjog pure vjjaiuavy Yi0q IeY) Os ‘(QERyT ‘UdIS19}SC sazorddyjog jo saisads-ad4}) aoy y, sunjou sndoysug yim IAIUIFUOD St (TOSI LOIUINIS sayzpsdquaydyjog Jo soisads-adA}) s1931n]g snzojaund sayrpsdquay -dyjog yey) uotuido ay) jo wie | Apsepiwtg -sn7 -pjaung ‘g yo AutXuouAs ay) ul (QF6]) SUIPIFY 5119048 Djjaluaivy IpNjout J ‘(T6I) [2494 2 191 -uoqisyD ‘osepodiyouy Aeyeyy ‘sourddiytyg ‘ — (49)1N|S) snynpIund sHojdtyjng ‘ot 0091-006 "y1OM siu) ‘(eggg6y) UOsmeg ‘eteIsny UJaYINOS pue UD)SBdYINOG ‘pueRjeaZ MON fsuvd9Q dYyloeg pue NUePY ul pesidsopi py ; _ (saeg) sunjou sajojddyjog °6 IMOY €Zb-26F "YOM SIU TL, — 3 sesuaypofiou (viunyjo1y9iuay) DiunyjosaPy §—°g sepnoeyyeuds Apiwiey 0£-0 “(G86T) 2Mow “(1261) MOY 2B 4121 foyloeg jesidos usa)seo¢ Sayloeg Isam-Opu] — (861) OUND 29 OquIODaSpy uossa] 0724 (visdso1mdy]) FY 02-0 “YOM SIU) SOyIIVg ISOM-OpUT — *; unsnsny tuzafop (nogoinnjs) Fy 9 £1-0 ‘(G861) Moy “(TZET) 2MOY 29 442ID Soyloeg jesidos) usojseog ‘oyloeg Isom-opu] — (¢861) NOUUDY 29 BquIODaSpy Jaduiag s1iiffip (nuosadtjyjg) ‘HG 01-0 (G86) MO’ “(TZ6T) MOU 7B HPI fouloeg jeoidos) us9a)seaZ ‘syle IsoMm-OpU] — (§ 861) NOUUDY 29 OqUIODaSpyY ss (Apueg) v70/2dso2Naq (vinyjorsuajsapy) “FY “F OI-0 “LO8I ‘eyUd[ag UOU feyxUD[Dg sz7opi0d (7) HS? “(G861) Mow “(TZ61) 2MOw BRIO soyploeg yestdosy usd)seag ‘syloeg Isom-OpuyT — (€ 861) WOUUDY 29 oquIODaSpy SIMPN’] Vjvaury (viunyjouossaT) FY 06-0 (C861) 2MOw “(1Z61) 2MON 41219 forloeg yesidos) usa)seag Soyloeg Jsam-Opuy - (€86]) NOUUDG 29 OquIODOSpy Jasoe[ myo (puapojvzyy) FY 7, 09§-ZF§ “YAOM SIU T, — " IMOY V29UN (DiunyjodaunA ) DIENY;O;OFT “| oeplunojoy Aprmey BOploinyIojOFY (uw) A691 oL91 “SofED0E < A691 oL91 ‘SoOEo8Z yidaq syUdWIWOD/A}ILIOYINY/UOIINGIIjSIGg 1324IO yueg eypouesue, puL[sy x[OJION uoxey, (psosas mau = ,) ‘saduns yidap puv suorjngiajsip jossuan yjim ‘vay uvUsD], UsayjsoU ‘yuDG DjjauDdUNAA puDv puvjsT yjofson WoL susapoutysa fo }5174234) T a1aV3L PROC. LINN. SOC. N.S.W., 111 (4), 1989 _ ee rr —- OnhvowmvwO—eeee ee esawsxXwx——eeeeee on 2 096-09 (9861) 1814s PID “1 'H ‘oseypodiysay Aejeyy ‘sourddiyiyg ‘uedef” — : 2p ZIssesy “\W snjojnavu snanajdoja0) 47, aepioeqiy Ajiweg CE-O8T “YAOM SIy) *(F6I) UOSUDILO ‘BITR.SNY ISVOD FN ‘spuels] .peuwoy ‘osepodiysay Aejeyy ‘sourddyiyg ‘uedef, — 5 ZISSeDY “Y wnsuo) vuapoipopidsp *¢z, osepnewoporpopidsy Aye 0S6-091 (9861) 9814S ‘osejodiyosy Aepey ‘sourddyiyg ‘uedef — A (zisseSy ‘y) 719048 vuosoiaajyy) °7z, 10§-O§1 “YOM SIT, - i IMOY wnsy7nd vuosojvgvzyy "|Z, aepiunyouryog Apwueg 0ZL-€8 (9861) !981yg ‘uedef = . (ulo}19poq) se719Aa suvpis0jdoyy °0% G9SZ-OLE (88961) SIU yO ‘purlesz mon] ‘ode YON ‘osepodiysiy Aepey ‘ues90@ ueIpuy] — (egg6]) YSU (zissedy “\) vuadysoy sizvprs0uajvg 6] 00S (9861) neaBOF ea) 2p MOY ‘(BLEMIYSOR ) DJDINILAgnjOLIIU “§ = 6 (G61) 1ysIUy IY = ‘ds stuppis0aajg gt iS 10£-69 (9861) noSS0H 2 omoy ‘(v9g ULUISE] ) ~ purjs] Moy ploy ‘osvjodyosy Aepeyy — (asaltayAy op) szppostaadg stavpr90jqjg * 1] cal (O€1é)€Z-0 (9861) NOBSOH 29 OmoY ‘(1261) MOY 2p = Y1P[D Spurjs] Ipeulsoy ‘oytoeg Is9m-OpuyT (GZ61) 1YsIUy I — (yoseure]) sypruadua snyjuvsvy yg 9 Ee (8614)68-O1 (9861) NOBSOH] 29 omoy ‘spurys] DopeUoyy ‘PURS] OMOF{ PsOTT ‘etpessny usd)seoy nog — (9861) 29580 F 2 OMOY NIBSOP 2 IMOY 7817/09 stuvpr90u0UT GT oepueply Aprmeg We] POM Ig 9¢-0 (961) PID “TH ‘(kag ueusez) pues] omoyY pso7y ‘eipessny usoyjnog = : (uosduing) vafizqvjop nidoudsojday “pI sepndeudg Aime 921 "YAOM SIU T, - — IMOY Wnpagiasod wnipiuotyjoany *¢] seproydoyAyg Apiurey “YOM SIU) ‘(GZ61) uosuR TY ‘epensny “seo UBLIO}IIA ‘BVIUBWISey “YN ‘osejodiyouy Aejeyy suede — ; ((99U,L) MjNIDU auoTOUjaDT “ZT (w) Ho69T- oL9I ‘Soko < Ho69I- oL 9 ‘Sof oz yidaq syUdUIWIOD/A}1IOYINY/UOTINGI414sIq 19419 yurg ejpouesuey pues] xJoy10N uoxey, (‘pauoD) I W1dVL PROC. LINN. SOC. N.S.W., 111 (4), 1989 NEW TASMAN SEA ECHINODERMS 284 OOT-0 GL-0 06-0 O0EC-8IE OSE-98I 06-0 00S-F6E 8LS-OLG 0S-O1 02-0 0£-0 (ur) widoq “YAOM SUA *(TZ61) MOY 2B ALTO ‘pueleoZ7 MON] ‘PULIS]T YON “}svoOo UOYIIOU pue spurs] 29peUlIoy ‘OyIoeg JSOM-OpuyT ‘(q¢961) uosmed :(1/61) MOY 7 AAL[D spuryeoz MON ‘PURIST YON “JSVOD ULOYIIOU ‘spueRys] JIPPULIOY Suipnyout ‘IIe JSOM-OPuUyT “(TL61) MOY 2B YARD oyloeg 1s9m-opuy] ‘(GZ61) 1YSIUyIW {(EF6]) UDsuUd OP] {spue]s] Iopeuoy Surpnyout ‘oyioeg-opuy ‘(¢461) uosudj10yy fosepodiyory Avjeyy ‘sourddriyg “(Injaqnop se pues] H[OJAON douRUdA0Id oy} PosopIis -U09 d}{ ‘sudutoads pjo om) Jo UOeUIUIeXd UO P1099 ST) posegq (¢P6]) UISUdIIOY] "YIOM SIU) SeIpessNYy Us1d}SeIyINOS ‘(GL61) JYSIUYIPY ‘puryeozZ Many ‘pueysy YON 10M Sty TL, "y1OM sty) “(7961) Jayeg ‘(eog ueusey) pues] MOF] Pso'7T *SITEAA YINOG MON] JO }SBOd ‘puR[eoZ MON] (TL6T) 9MOY 2B YAEL syle Jsom-opuy (46961) uosmeg {puerleoZ MON ‘pue|s] YON JO JsvOo uLOYyIIOU ‘spues] Iopewioy {(eog ueUIsSey ) puryjs] Mop pso07 feipessny usojseoy nos syusuIW0D/ALIOYINY/UOINGII4sSIG 19410 Ao69T- oL91 ‘SofEo0E< = (¢861) NoUUDg 2 squIoD9Spy = (€861) HoUUDY 2 equIoDD5py * (GZ61) 148IU yO = (E461) Uosud OW = é (GL61) ystUy IN — * = (€96]) NoUUDg 29 aquIoDDSpy A 069I- oL9I ‘So0Eo8Z yueg ejpoursuey puvy[s] Y[OJION (UIpPDYDIJA]) VuDIpUr DYajogopnasy GE (snowuury) 7727010 sajsnauduy "HE (youeure’y]) snjoajd sajsnaudoxoy —¢¢ sepnsnoudoxoy Ajriwmey (zissedy “W) 2uijouom snurysug *ZE UISUIJIOPY pLodosvu SLDpr9z0uOs IL], TE ZISSeSY UL UdyIN'T] snpojfur sajsnaudojoyy QE sepiinojdouwnay Apiwmey UOSMEY aDIpUD]DazavAOU DuIpadouavy GZ, IMO LayDquonjD Duipagouan’) 8S seprurpog Ajrumeg JIayeVg waupod vuapyiq [7 UL[OYIIJY -2dusiavs DuapoiqT OZ (zissesy ‘W) 2suadpor snuvydajsouay GZ, sepnewoperg Ajmey uOoxey, (pauoD) | aTdVvL PROC. LINN. SOC. N.S.W., 111 (4), 1989 285 OZI-0 (L961) OAeY :purleoZ MON “pur|s] YON ‘svoo uLoYIIOU ‘(og UBUUSR],) PUPS] IMOP{ PLOT ‘eIpeaysNy Isvoo uso}seo Suede’ _ (£961) 299% g ULIOpOG 1zssvsn snssug “OF depissiig Ajiwiey Ibl-GE (G96) Uosmeg ‘(e9g UR -Se],) pues] MMOFY PsOT soloed SoM-Opuy — (qG96]) Uosmeg (sua}1e Jy UOA) snuodrda mipodos1¢C Gy sepisAdojedy Ajrwmey 09%-L1 (88961) }YSIUYpPI ‘Spue]s] Ipeulioy ‘purjeozZ MON (89961) 1YsIUy II (qg96]) Uosmeg UDSUDIOJY aVowaury DaUoLayT “HH 10€-S (IL61) MOY 2B YAPTO sosepodryosy Aeyeyy ‘sourddiyiyg ‘ues0Q uRIpu] Usd)}seq — i (ay[taurelg) yvuodn2ap wnunsvyT Sy aepiuesey Apiwmey 9€C-6 ‘(qGg6]) UOsMeg ‘puRlesZ MON ‘pue]s] YON — qcgg6] ‘Uosmeg UISUI}IOYY SnL0gdjog snumdzouyIy “Zh oepraeynq ty Apres 1O€-OF (9861) 9814S ‘puryeoZ MoN pure elpeaysny ‘eury-opuy ‘sourddiyryg ‘uede[” — A UID[IapO) suarsaua saysvaddq]y “|h = 0G-¢ ‘(qG96]) UOsmeg ‘spurs] 997peUtisy é ro) {puerjeoZ Man] ‘pur|s] YON “sBO09 (a4 usJoysou ‘(vag URWUSe]) puL[s] MOY PLOT ical ‘elpersny usojseq ‘uedef ‘oyioeg jsamyyNog (GL61) 1UBIUSPIIA qGQ6] ‘UOsSmeg (Aeris) avisvjoujsnv sajsvagay) Or = : sepliaqseaddy Apiurey foo 0€-0 ‘(TL61) MOY 2 YARD ‘oyloeg jsom-opu] — (861) HOUND 29 oquIODASpY (snaeuury) snyopprumuou snjoyuarolajaFy 6 $S-0 (46961) UOSMEY {PUR]|RIZ MON ‘PURIST YON “Iso UJoY OU ‘spuRIs] IpeULIoy ‘(wg UBUISP] ) purys] oMoP] Psoy Serpessny uso}seoyINOG = (€86]) NOUN 2 OquIO.aSpY (yorewie'{) vivjns1aqgn} suuppr90NaFT «BE 6£1-0 (4961) ¥osmed “(TL61) IMOY 29 FAL] ‘SPUL]S] DIPBWIIY ‘vog ULI -SB], ‘(PUR]S] OMOP{ POT 9yI9eg JSOM-Opu] _ (€g6]) HOUND 29 oquIODaSpy (a]ptaurelg) zavyjow vijamouryrg “LE seplujowoulyog Ape FVCEI-OIG YAOM STY} ‘IPGRQOS] “IYSIUYpy ‘eIpessny uUsO}Seoy Nos {pueleozZ MON ‘spuey]s] Iopewoy (GL61) 1YSIUyPI = (44RD TH) smapjuaprynwu snurysajvmy 9 oepruryog Apruey (ur) F691 oL91 ‘Sokfo0E < F691 oL91 ‘So0Go8z yidaq syUIWIUIOD/A}IIOYINY/UOTNGII4SIg 1924319 yurg epjoursue puL][s] Y[OJION uOXxey], A ee ee ee en tree pe el a nr gy RR el ee ee ee SS (‘pauoD) | a1dVL PROC. LINN. SOC. N.S.W., 111 (4), 1989 NEW TASMAN SEA ECHINODERMS 286 BEI = eQ96I YSU SIN = inuviyj vanryqorygup 6G 9EZI- 406 (6261) jayeg ‘vag wiesdD ‘sourddiryg ‘iwemepy — (6261) 294k g (49] Yoox) V7270su2 viniydoryqup “gc 806-012 (6261) 24a ‘SPULIS] IPPULIOY -jSVOO URI[eIISNYy U19}SvO ‘osejodiyouy Aeyeyy ‘sourddipiyg ‘uede(” — 5 WRID TH vyjunsviniw pimiydg "LG sepriniyd Aprumey 806-SE ‘(Ig61) ong