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Cee eet Co eae OMe iL SL eae s SOU SrT re CUR i ia ed Wye ne pees Spb ie’ AUT easy CESS pret oe peda AOD Ey Gye ieee wan See yor cee® TE ey, hg eo Weererr at teeny rely A oye hie! ees seluyuaes ate 4 eae Pere es ny eee vy POP Oe ae een Se wee par wee na yee Re eponees Gage eae sayy wie si3y Sy gageycaeweayaqepe vey at Penk pipaigns Cdk et sare wh ep Cae NT Ava eh ERE L weve Peta Ais APES are te ale dehd syd ote ay Perna Tet Ta MaMa ea ele ’ pia dan eget BSS hearts Lo Leas AHL Cote Wipe (ERS Gf AO 84s Ras Fe 2 Ee anya R ia EAA DD eS 1 iy eae A yiga 2d Ae td Reg reese NUTS Be Peere ess waa ge as To Peay Bait dU ANDREA o wee Oa crete Haeautp ype vena an ye aawsh e bee we eas a a ee a al ae ao vert heb od PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 110 NUMBER 1 NUMBER 2 trees NATURAL HISTORY IN ALL ITS BRANCHES fe EINNEAN S@OCiInIY Or 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 1s $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 1986-87 President: P. M. MARTIN Vice-presidents:)s HELENE A. MARTIN, G. R. PHIPPS, C. N. SMITHERS, A. J. T. WRIGHT Honorary Treasurer; A. RITCHIE Secretary) BARBARA J. STODDARD Council: A. E. J. ANDREWS, M. ARCHER, JUDITH HH. K. EASIPMAN, Ra FAGER, L. W. GC. FILEWOOD, MARILYN D: FOX, EL. A: S; JOHNSON; HELENE VAD MARTIN (EMC OMARIING ) J, Re IVEIRIR TOK Gaia. MYER SCOUGH 1) G.oPERCIVAL: GAR) RHIRPS) Ay RiGC Hib Case SMITHERS, T. G. VALLANCE, KAREN L. WILSON, A. J. T. WRIGHT Honorary Editor: T. G. VALLANCE — Department of Geology & Geophysics, University of Sydney, Australia, 2006. Linnean Macleay Fellow: R. W. JOHNSTONE Linnean Macleay Lecturer in Microbiology: K.Y. CHO Auditors: W. SINCLAIR & CO. The postal address of the Society is P.O. Box 457, Milson’s Point 2061, N.S.W., Australia. Telephone (02) 929 0253. © Linnean Society of New South Wales Cover motif: ‘Fossils of Talbragar’, a vignette designed and drawn for this volume by Alan E. J. Andrews. PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES Marine Biological ‘Laboratory LIBRARY Dec 21988 =| Woods Hole, Mass. VOLUME 110 NUMBER 1 Diurnal Survey of Ichthyoplankton Abundance, Distribution and Seasonality in Botany Bay, ; New South Wales ALDO S. STEFFE and BRUCE C. PEASE STEFFE, A. S., & PEASE, B. C. Diurnal survey of ichthyoplankton abundance, distribu- tion and seasonality in Botany Bay, New South Wales. Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 1-10. We found 37 types of pelagic eggs, and 27 taxa of fish larvae representing 12 families, during an ichthyoplankton survey in Botany Bay from September 1977 to October 1979. A small plankton net was used to take surface samples during the day. Pelagic eggs were most abundant during February (late summer) of both years. The abundance of pelagic eggs decreased significantly with increasing distance into the estuary from the bay entrance. The greatest number of larval fish taxa was captured in March (early autumn) of year 1. Although Botany Bay is physically best termed an ocean embayment, its larval ichthyofauna is similar to that of other temperate estuaries worldwide, in that larvae derived from demersal eggs dominated the assemblage. Larvae hatched from demersal eggs contributed more than 78% of all larvae collected. The percentage of larvae derived from demersal eggs at each station increased signifi- cantly with increasing distance into the estuary. Aldo S. Steffe, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109, and Bruce C. Pease, Washington Department of Fisheries, Point Whitney Shellfish Laboratory, PO. Box 102, Brinnon, WA 98320, U.S.A.; both formerly Fisheries Research Institute, N.S.W. Department of Agriculture, Cronulla, Australia 2230; manuscript received 19 February 1986, accepted for publication 22 October 1986. KEY WORDS: Ichthyoplankton, fish larvae, pelagic eggs, distribution, seasonality, Botany Bay, temperate estuary, ocean embayment. INTRODUCTION The only published study of ichthyoplankton in an Australian estuary is that relating to the Blackwood River estuary in Western Australia (Lenanton, 1977). Eggs and larval stages of some Australian marine fish species have been described (Tosh, 1902, 1903; Dakin and Colefax, 1934, 1940; Munro, 1942, 1945, 1948, 1953, 1955; Thomson and Bennett, 1953; Cassie, 1956; Thomson, 1963; Robertson, 1973, 1975; James, 1976; Leis and Rennis, 1983) but there is still little Australian ichthyological literature in the fields of egg and larval development, and estuarine ichthyoplankton ecology. Our object was to describe the ichthyoplankton assemblage of Botany Bay, New South Wales, and to compare the findings with those from other temperate estuaries worldwide. MATERIALS AND METHODS Study Area Botany Bay (34°01’S, 151°11’E) is a large, semi-landlocked estuary (Fig. 1 ). The bay has a surface area of 4160ha and receives discharge from the Georges and Cooks Rivers. Despite this riverine influence the bay is dominated by ocean swell and wind waves (Roy et al., 1980), is vertically well mixed (Rochford, 1951), and at most times is best described as a marine embayment. The bay has extensive areas of seagrasses and mangroves. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 2 BOTANY BAY ICHTHYOPLANKTON Definitions Ichthyoplankton: portion of fish community found in the plankton and which have not attained juvenile status. Larval assemblage: all larvae caught within the constraints of the sampling programme. That is, by daytime sampling within three hours of high tide. The terms pelagic egg, demersal egg, larva, and juvenile all follow the definitions of Leis and Rennis (1983). In Botany Bay the families Gobiidae, Blenniidae and Syngnathidae which 1s ovoviviparous were considered to have demersal eggs, and all other taxa were regarded as having pelagic eggs. Year 1: September 1977 to August 1978 inclusive. Year 2: September 1978 to August 1979 inclusive. Taxon diversity: number of taxa present at all stations during a given month. Taxa identified to family level are only regarded as a single unit. Taxon diversity was not cal- culated when only unidentifiable yolk-sac and damaged larvae were represented that month. Collection of Samples Plankton samples were collected at nine stations each month from September 1977 to May 1978 then from August 1978 every second month to October 1979 (Fig. 1). A further station, Mangrove Channel (MC, Fig. 1), was sampled only until October 1978. A 27.5cm diameter net with 350um mesh was used for the monthly samples to May 1978 when it was lost. A net with mouth diameter of 40cm and the same mesh size was used thereafter. The 40cm diameter net which filtered about twice as much water as the first net was used for the rest of the study. All ichthyoplankton counts from samples col- lected with the 27.5cm diameter net were doubled to account for the increased filtering capacity of the second net. At each station two replicate samples were collected. All sampling was carried out around high tide (+ 3hrs) during daylight. The plankton net was towed for five minutes (+ 5secs) at the surface at a speed of approximately 1.0m sec!. Although a flowmeter was not used to quantify the amount of water filtered the towing speed for each sample was kept the same by setting the number of engine revolutions at a constant value. Immediately after collection all plankton samples were preserved in 10% buffered for- malin. All samples were sorted entirely, using a dissecting microscope. Larvae were identified using the size series method (Leis and Rennis, 1983) and pelagic egg types were identified using the characters outlined by Robertson (1975). Equipment failures and inclement weather sometimes prevented completion of the full sampling programme for a given month. No stations were sampled during June 1978. Stations not sampled at other times were the Northern Bay (September 1977), Mangrove Channel (March 1978), Towra Point (August 1978), Woolooware Bay and Georges River (October 1978), and North-Central Bay (April 1979). RESULTS Limitations of Data Set The limitations of the data set this manuscript is based on can be summarized as follows: 1. Individual sample sizes, i.e. the volume of water filtered per tow, appear to be too small. In order to rectify this we decided to pool samples. Pooling enabled us to increase sample sizes to an acceptable level but decreased the available degrees of freedom for PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 ALDOS. STEFFE AND BRUCE C, PEASE 3 Fig. 1. Location of sampling stations in Botany Bay. Abbreviations are: PE = Port Entrance, PA = Port Area, NC = North-Central Bay, QB = Quibray Bay, NW = North-West Bay, TP = Towra Point, NB = Northern Bay, WB = Woolooware Bay, GR = Georges River, MC = Mangrove Channel PROC. LINN. SOG. N.S.W., 110 (1), (1987) 1988 4 BOTANY BAY ICHTHYOPLANKTON statistical tests. Therefore the trends we observed must have been strong because the fewer samples the more marked a trend must be to give a significant result. 2. Larval avoidance of the small nets was potentially large. We resolved this dilemma by restricting our survey to pelagic eggs and early larval stages which our nets sampled adequately. 3. Although we acknowledge the larval assemblage would have been better estimated using larger nets, and by sampling at night we believe our samples represent accurately the relative proportions of early stage larvae hatched from demersal, and pelagic eggs present in the larval assemblage. Species Composition The pelagic eggs collected were found to comprise 37 types, however, only the oval egg of the anchovy Engraults australis could be identified with certainty. Twenty-seven taxa of larvae from 12 families were collected (Table 1). The bulk of the larval assemblage were species from demersal eggs (78.9%), with Gobiid and Blennid larvae being the most abundant. Spatial Distribution of Ichthyoplankton Pelagic eggs were most abundant at the two stations nearest the entrance with the Port Entrance and Port Area stations (Fig. 1) accounting for over 76% of all eggs col- lected (Table 2). The abundance of pelagic eggs (logarithmically transformed) decreased significantly with increasing distance into the estuary from the Bay entrance (Fig. 2a). Larvae from demersal eggs were dominant in the catch at all sites except the Port Entrance. The percentage of larvae from demersal eggs at each station increased signifi- cantly with increasing distance into the estuary (Fig. 2b). Seasonal Abundance In both Year 1 and Year 2 the peak in abundance of pelagic eggs occurred in February (Fig. 3). Larval fish catches in excess of 40 were recorded during October, November, February and March of Year 1; and February of Year 2 (Table 1). The October peak in Year | is due to a catch of 108 individuals of goby species 4 at Woolooware Bay. Larval fish abundances in Year 1 peaked during October-November (Spring) and February-March (late Summer-early Autumn), whilst during Year 2 only the February (late Summer) peak was repeated (‘Table 1). Taxa Diversity of Larvae Four taxa of larvae or more were caught in October, November (Spring) and January, February, and March (late Summer — early Autumn) of Year 1 and Decem- ber, February, April (Summer Autumn), and August (Winter) of Year 2 (Table 1). Taxa diversity was low at other times. The diversity of fish larvae we found was extremely low compared with the total of 229 species of fish recorded from Botany Bay during the same period (Anon, 1981). DISCUSSION Whilst we acknowledge that the data set is limited due to some problems associated with the collection procedures (see Results) we believe that this survey is representative of the pelagic egg and early stage larval component of the ichthyoplankton. We interpret the large proportion of pelagic eggs found near the bay’s entrance and the observed PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 ALDOS. STEFFE AND BRUCE C. PEASE G G I 9 9 4 I 0 G I O} 9 G 6 9) L G L I 9 Gi LI 6 I v G COT 69 96 OT bh 8cl oa = = = 6 re a I = = OT re (6 = =e se = G Es I I i = ce es ea t G = 6 ee 9 I =F if == = = a a ia = t G = I LG a 6 = me a 16 96 ¥G t 6G BIT — — — _ — — — — G = = — — = =e ae I OT G =a i a = all 6G 9 v 91 za = es = safe aa = = 6 — —_— — = = — G — — —_— — —_— = I vE ae v — _— _— = — a = G — — = sa aa oS a ce = ee = ne = 6 if — _— _— _— — — — — — — — — — — — — = = — — = = — & —_— = = I I = = = = = = t = = = 6 = < = = I G = a = = 7: ae a ry 5h = = C = = 6 6 —. = = — G — = — pO sny ounf ady qoay 92q WO sny AeypFw ady sme gqoqy uel 2q AON WO 6L61 8Lol LLO6I (paj00d SU017D79' 7] VY) IDQLD'T ysiy {0 Cassaarg UOXDT Puvd IJUDPUNG Pp JDUOSDAS: T dav Lb AVISIOATC] UOX® J, RAIL] “ON [BIO], BAIL] paseurep pue oes-yJOA aIqeynusptuy (dds 7) exe] umouyug “ds nruayasnapy oepryluPRoeUuoyy (dds 4) sepigon sndny sapaasoLjag SNIJOUD SNYIUDLGOWC) sepluusl”_ DIJUITLD VZVT oeprsnyl pyoprdsn9Ir4) vp) a4) aepisoydAy snajuasin snjdjavpouop aeplAioepouopy aepuedg snjoao sassay) aeplot4wayy DIY) OBDINS DePIUIBLTIIS SUSI] DIZ0X SLAJSOLIULLDI SNGULDIOL afoul snyyousudy DAdIU DLOGO]DULBI]S SNB1D DLOGoO;DULB IIS aepiyieusudg seplulloyyy seprydoA yy (1ap10 DTUTOUOXe} ul posursie) VXVL PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 6 BOTANY BAY ICHTHYOPLANKTON WABLE,2 Total Abundance of Ichthyoplankton and Percentage of Larvae from Demersal Eggs at Each Station Distance from No. Pelagic No. % Larvae from Station entrance (kms) Eggs Larvae Demersal Eggs Port Entrance PES) 1472 13 38.5 Port Area 4.0 874 24 66.7 North-Central Bay 5.9 208 18 66.7 Quibray Bay Fell 123 14 85.7 North-West Bay Was 51 56 58.9 Towra Point 7.5 155 113 85.8 Northern Bay 7.6 147 oA: 63.0 Woolooware Bay 11.1 we 153 94.8 Georges River IBIS: 27 31 83.9 Mangrove Channel* 6.1 4 8 25.0 *Station Excluded from Regression Analyses as sampling was discontinued after October 1978. decline in numbers further into the estuary (p<0.001) to be due to most species with pelagic eggs spawning near the bav entrance or outside the estuary (Bell, 1980), and, the subsequent ‘dilution’ of pelagic eggs within the estuary by localized tidal and wind currents. Although Botany Bay is characterized by its marine qualities (Roy et al., 1980) it has a larval assemblage dominated by taxa of estuarine origin, 1.e. larvae hatched from demersal eggs. No apparent trend was found between distance from the bay entrance and the abundances of larvae derived from pelagic eggs, or those hatched from demersal eggs. However, there was a change in the relative proportions of these larvae with increasing distance into the estuary. The percentage of larvae derived from demersal eggs at each station increased with distance into the estuary (p <0.02). This trend is not surprising as most species with pelagic eggs spawn outside the bay or near the entrance (Bell, 1980), and many species with demersal eggs spawn within Botany Bay (Bell et al. 1984, Middleton et al. 1984). Because larvae from demersal eggs hatch at an advanced stage of development, with a better swimming capability than comparable larvae from pelagic eggs they are more likely to be retained within the estuary. Thus the earlier stages of larvae hatched from pelagic eggs could be present in Botany Bay only in low numbers, with active migration into the estuary probably occurring later, at larger sizes which were capable of avoiding our small nets. The February peaks in abundance of pelagic eggs correspond with the late Summer peak in spawning of economically valuable fish species found by Bell (1980) in Botany Bay. The peak of pelagic eggs in February of Years 1 and 2, the larval species diversity peak in March of Year 1, anda subsequent large catch of larvae in mid-March 1981 (Steffe, 1982) suggest that the peak spawning period of most fish (in- cluding those of no economic importance) in the bay occurs in late Summer (February) — early Autumn (March). Whilst Botany Bay has been shown to be an important transitory nursery area for many commercially valuable fish species (Bell, 1980; Anon, 1981; Bell et al., 1984; Middleton et al., 1984) it appears that relatively few of these species utilize Botany Bay during all phases of their life cycle. Although the Botany Bay estuary is, physically, best termed an oceanic embay- ment, its larval assemblage closely resembles the estuarine larval ichthyofaunas of other temperate regions. In Botany Bay, larvae hatched from demersal eggs constituted 78.9% of the larval ichthyofauna with the Gobiidae being the most abundant taxa. Similarly, the larval assemblages of other temperate estuaries in Western Australia, PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 | ALDO S. STEFFE AND BRUCE C. PEASE ° 3 => 1000 3 o 100 8 A 10 r2=0:911 p<0-001 o ¥=3-62 —0:204x 2 2 4 6 8 10 12 Distance from entrance (km) 100 eggs 80 r2=0-551 p The three species G. brimblecombet, G. blakec Moore and G. eremica Moore all utilize NEEP E. camaldulensis as host, in three areas each of which overlap in part, throughout the host’s extensive continental distribution (Moore, 1971, 1975, 1978). This may indi- cate that the eucalypt species consists of three separate taxa, though two only are at present recognized. K. In a suggested general phylogeny of the eucalypt ‘subgenera and suballiances (Johnson, 1972: 14; 1976: 159), the Monocalyptus suballiance is placed before the Sym- phyomyrtus suballiance, whereas Monocalyptus is placed after Symphyomyrtus in Pryor and Johnson, 1981: 504. It appears that the species of subgenus Glycaspis (Synglycaspis) constitute the most ancient group in the genus because of the number of antennal rhinaria and their dis- position (Eastop, 1958), and because of the gall-forming, flat and round-lerp forming habits of the species (Moore, 1961 et seq.). Species of the subgenus Glycaspis (Glycaspis) construct the more advanced lerp shapes of round, oval, cloverleaf, square and rectangular, and bear progressively fewer antennal rhinaria, thus indicating that the group is of more recent origin than Glycaspis (Synglycaspis) in the evolutionary sequence of the genus. The Symphyomyrtus suballiance may thus be considered of more recent origin than the Monocalyptus suballiance. L. The species G. atkinsoni Moore (1984) colonizes EFAAA E. similis Maiden, and appears to be nearest to G. violae Moore (1970) which is known to occur on SUG:A E. cambageana Maiden and possibly SUP:V E. melanophloia F. Muell. E. cambageana is included in the ‘box’ group of eucalypts, and EF. melanophloia in the ironbark’ group (Chippendale, 1968). E. similis is included in the ‘subgenus’ Eudesmia of Pryor and Johnson, 1971. It is suggested that E. szmilis may have affinities with the ‘box’ and ‘ironbark’ groups of Eucalyptus. HOST SPECIFICITY Indications now suggest that certain species of Glycaspis are host-specific, and others may be termed Series-specific in the context of Pryor and Johnson’s classification of the eucalypts (1971). The above two categories of Glycaspis are listed, with the hosts alphabetical. Such lists are obviously incomplete and prone to revision because of limitations on past studies. They will provide a ready reference to most Glycaspis species hosts, thus facilitat- ing identifications of Glycaspis species. Lerp shapes are referred to in the lists by the following letters: G = galls; F = flat lerps; R = round; R-O = round to oval; CL = cloverleaf; S = square: RT = rectangu- lar lerps. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 22 ASSOCIATIONS OF SOME GLYCASPIS SPECIES Glycaspis spp. regarded as host-specific. Subgenus Synglycaspis ampluficata on E. acmenoides G encystis on agelomerata G brunosa on coccifera G surculina on conglomerata G wallumaris on conglomerata R morgant on diverstfolia G ecphymata on dives G perthecata on haemastoma G wcterica on marginata R munita on nitida G phreata on oblonga Ie belua on pauciflora G cyta on pilularcs G sertata on pilularis R cyrtoma on piperita G planaria on piperita F longaeva on pulchella G nundlensis on radwata F immaceria on rOSS1L G tagmata on rodwayt 18 obvelata on ?svebert G constricta on stricta G inclusa on umbra G orientalis on umbra R Subgenus Glycaspis riguensts on . argophlora R-O inusitata on bakert S, RT rylstonensis on blakelyt oR clivosa on brachyandra R rubritincta on brevifolia R onychis on brevifolia R blake * on camaldulensis CL eremica on camaldulensis CL emphanes on cambageana Ral subita on cornuta R deirada on dundasit RT montana on dunnit R exsertae on exserta R keremae on exserta R-O monita on fasciculosa R-O flavilabris on gontocalyx R caurina on jensen R infucata on leptopoda R collina on melanophloia R *one authentic record on ferelicornis PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 neureta egregia mannifera praescopula lacustris pilata Kurrajongensis mellialata permista suavis umponens operta stliceflava xanthopepla sudicola atkinsont nancyana struicis felicitarts lucrosa wagaityae dobsont on on on on on on on on on on on on on on on on on on on on on on K. M. MOORE melliodora R-O moluccana R moluccana R orgadophila R-O ovata R-O paniculata R paniculata R-O paniculata RE@® paniculata R-O populnea R punctata R raveretiana R robusta RT seeana R sideroxylon R-O similis R squamosa R tereticornis R tetraptera R tetrodonta R ? tetrodonta R viminalis R Glycaspis spp. regarded as Series-specific Subgenus Synglycaspis aggregata on 2 spp. conserta on 2 spp. cyanoreia on 2 spp. endasa on 2 spp. salebrosa on 2 spp. temenicola on 2 spp. Subgenus Glycaspis wondjinae anota quornensts aurosala whiter baileyr wgnea granulata johnsont pratensis minuscula BLAKELY, W. F., 1955. — A key to the eucalypts. 2nd ed. Canberra: Forestry and Timber Bureau. CHIPPENDALE, G. M., 1968. — Eucalyptus buds and fruits. Canberra: Forestry and Timber Bureau. on 3 spp. on 2 spp. on 2 spp. on 2 spp. on 2 spp. on 3 spp. on 3 spp. on 3 spp. on 3 spp. on 2 spp. on 4 spp. in Series Haemastominae in Obliquae in Obliquae in Piperitae in Piperitae in Piperitae inSeries Albae in Miniatae in Moluccanae in Ochrophloiae in Polyanthemae in Salignae in Salignae in Salignae in ‘Tereticornes in ‘Tereticornes in Viminales References AA AAAA T I R-O (ETE 23 PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 Mee ASSOCIATIONS OF SOME GLYCASPIS SPECIES Eastop, V. F., 1958. — Some neglected taxonomic characters of Psyllidae (Homoptera). Ent. Mon. Mag. 94 (1124): 18-19. JOHNSON, L. A. S., 1972. — Evolution and classification in Eucalyptus. Proc. Linn. Soc. N.S.W. 97: 11-29. ——, 1976. — Problems of species and genera in Eucalyptus (Myrtaceae). Plant Syst. Evol. 125: 155-167. Moore, K. M., 1961a. — The significance of the Glycaspis species associations with their Eucalyptus spp. hosts; erection of a new subgenus and descriptions of thirty-eight new species of Glycaspis. Proc. Linn. Soc. N.S.W. 86: 128-167. ——.,, 1961b. — The biology and occurrence of Glycaspis baileyi Moore in New South Wales. Proc. Linn. Soc. N.S.W. 86: 185-200. —, 1970. — A revision of the genus Glycaspis with descriptions of seventy-three new species. Aust. Zoologist 15: 248-342. —— , 1971. — The Glycaspis spp. — Eucalyptus camaldulensis associations. J. Ent. Soc. Aust. (N.S.W.) 7; 3-7. ——., 1975. — The Glycaspis spp. associated with Eucalyptus camaldulensis. Proc. Linn. Soc. N.S.W. 99: 122-128: —., 1977. — Two new species of Glycaspis Taylor from Western Australia. J. Aust. ent. Soc. 16: 253-255. —, 1978. — Further information on Glycaspis species associated with Eucalyptus camaldulensis, and other Glycaspis species. J. Aust. ent. Soc. 17: 257-260. ——, 1983. — New species and records of Glycaspis Taylor with phyletic groupings. /. Aust. ent. Soc. 22: 177-184. ——, 1984. — Two new species of Glycaspis from tropical Queensland, with notes on the genus. Proc. Linn. Soc. N.S.W. 107: 475-478. —., 1985. — Four new species of Glycaspis Taylor (Homoptera: Spondyliaspididae) from some endangered species of Eucalyptus. Proc. Linn. Soc. N.S.W. 108: 71-78. Pryor, L. D., and JOHNSON, L. A. S., 1971. — A classification of the eucalypts. Canberra: Aust. National University. , and ,, 1981. — Eucalyptus, the universal Australian. In A. KEAST, ed., Ecological biogeography of Australia. The Hague: W. Junk. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 A new Species of Glycaspis (Homoptera: Spondyliaspididae) and some new Host Records K. M. MOORE Moore, K. M. A new species of Glycaspis (Homoptera: Spondylhiaspididae) and some new host records. Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 25-26. A new species of Glycaspis Taylor from Collinsville, Queensland, is described and figured, and some new host associations and distributions are recorded. K. M. Moore, Statue Bay, Yeppoon, Australia 4703; manuscript received 3 June 1986, accepted for publication 18 February 1987. INTRODUCTION With the description of this species, the subgenus Glycaspis of Glycaspis Taylor 1960 now consists of 88 species which are confined to the Symphyomyrtus suballhiance of Eucalyptus UHerit. (Pryor and Johnson 1971). DESCRIPTION AND NOTES Glycaspis (Glycaspis) incomperta sp.n. Figs 1-3 Types. Holotype ©’ on slide labelled Collinsville, Qld, 22 111 1984, A. G. Webb, P052. Paratypes, 2 slides of single wholemount males with same label data, but PO93 and P094 respectively. Claspers and aedeagus as in Figs 1-3. Length of aedeagus 0.195-0.205mm (3 specimens). All specimens are 1n the Australian National Insect Collection, CSIRO, Canberra. Notes. The phyletic position of G. incomperta is in the taylor: group, and appears to be nearest to G. xanthopepla Moore from which it differs in the following characters: the aedeagus is longer, with its distal end wider and more curved; posterior border of claspers more rounded, distal expansion of less area, the basal ‘foot’ of broader profile, fewer setae on external face. The host of G. xanthopepla is Eucalyptus seeana Maiden the northern distribution of which reaches to about latitude 26°S. Collinsville is about 700km in a direct line from the nearest known occurrence of E. seeana. Etymology: Latin, incomperta = of which one has no information (on lerp shape or host). NEW HOST RECORDS Glycaspis (Borevoglycaspis) devexa Moore has now been found to colonize Melaleuca dealbata S. T. Blake, at 10km S Yeppoon. G. (B.) australiensis Moore occurs on Lophostemon (= Tristania) suaveolens (Soland. ex Gaertn.) Wilson and Waterhouse, at Byfield State Forest, Yeppoon. G. (G.) brimblecomber Moore colonizes Eucalyptus mannifera Mudie ssp. maculosa (R. 'T- Bak.) L. Johnson, and E. brassiana S. 'T. Blake grown from seed. Both hosts occurred at Canberra, A.C.T. (M. Carver, pers. comm., 1986). _ PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 26 A NEW SPECIES OF GLYCASPIS Figs 1-3. Glycaspis incomperta sp.n. 1, aedeagus; 2, clasper, internal face; 3, clasper, external face. Posterior border of claspers on left. Scale line 0.1mm. G. (G.) minuscula Moore previously known from E. cinerea F. Muell., E. rubtda Deane and Maiden and E. ?viminalis Labill., also colonizes E. pulverulenta Sims at Cox’s River on the Lithgow — Jenolan Road, N.S.W. (Collections made by K. G. Campbell, 1987). References Pryor, L. D., and JOHNSON, L. A. S., 1971. — A classification of the eucalypts. Canberra: Aust. National University. TayLor, K. L., 1960. — Additional information on the Australian genera of the family Psyllidae (Hemi- ptera: Homoptera). Aust. J. Zool. 8: 383-391. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 First Report of Late Devonian ‘Trilobites from eastern Australia A. J. WRIGHT WRIGHT, A. J. First report of Late Devonian trilobites from eastern Australia. Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 27-30. ‘Iwo generically indeterminate specimens of a phacopinid trilobite, of probable tolV — VI (late Famennian = late Late Devonian) age, are described and illustrated. From the Mandowa Mudstone near the township of Barraba, New England, N.S.W., these are the first recorded Late Devonian trilobites from eastern Australia. A. J. Wright, Department of Geology, University of Wollongong, PO. Box 1144, Wollongong, Australia 2500; manuscript received 22 April 1987, accepted for publication 17 June 1987. INTRODUCTION Some years ago two trilobite specimens were collected by John Irving from mudstones outcropping adjacent to the Manilla River, east of Barraba, N.S.W. (Fig. 1). Both specimens lack any cephalic material (Fig. 2) but are clearly phacopid in nature, and are believed to be members of the subfamily Phacopinae as shown by the nature of the pygidium. They are recorded in this note as they are the only known Late Devonian trilobites from eastern Australia. AGE AND STRATIGRAPHIC ASSIGNMENT The locality (Fig. 1) falls within an area attributed by Chesnut et al. (1973) to the Late Devonian Mandowa Mudstone. Outcrops east of the town of Barraba (GR 278354) contain abundant Leptophloeum australe and rare shelly fossils including the trilo- bites recorded here, and may well have formed the original basis for the ‘Barraba Series’ (Benson, 1913). Mory (1982, fig. 4) gave an age of Early Carboniferous for the Mandowa trilobite localit a y Fig. 1. Sketch map of Barraba township, showing approximate locality from which specimens were collected. Australian Map Grid, spacing ikm. Taken from Piedmont (9037-III-N) and Barraba (9037-TII-S) 1:25,000 sheets. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 28 LATE DEVONIAN TRILOBITES LTE E 8 S LEE LEEDS beoomron ha ae Mia 1 PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 A. J. WRIGHT 29 Mudstone at Barraba, on the basis of the conodont Szphonodella quadruplicata. Dr Mory informs me (pers. comm., 1986) that his material was from outcrops west of the bridge on the northern side of the town, and that this locality appears to be considerably higher stratigraphically than the trilobite locality. Mory (1982) summarized conodont and ammonite data from the Mandowa Mudstone (incorporating data from Pickett, 1960; Jenkins, 1969, zn Roberts, 1972), showing that fossils as old as the Platyclymenia — Stufe (tolV) of the Famennian (= upper part of Late Devonian) are present, and that conodonts from other localities suggest ages as young as In3 (Tournaisian = Early Car- boniferous). The lycopod Leptophloeum australe occurs on the same rock as one of the trilo- bite specimens and is highly suggestive of a pre-Carboniferous age (Dr N. Morris, pers. comm., 1985). Genera to which the trilobite specimens might belong on morphological grounds are: Cryphops (tol-IV); Dianops (tolV-V1); Neophranops (toI-II1); or Tremerocephalus (tol- III). Given in parentheses are the ranges of genera as stated by Richter and Richter (1955, fig. 1) and Hahn and Hahn (1975) in terms of the German to (= Stufe or stage) ter- minology. (These subdivisions of the Late Devonian [see e.g. Roberts et al., 1972, chart] are approximately tol = Frasnian and toII-VI = Famennian). These data indicate a Late Devonian age for the Mandowa Mudstone. However, on the basis of Mory’s (1982) discussion, a maximum age of tolV is indicated for the Mandowa Mudstone. As only Cryphops and Dianops range above tollIl, the specimens may be favoured to belong to one of these two genera or perhaps an undescribed genus. Identification to genus is not possible in the absence of cephalic material. Assignment of the material to the Phacopinae 1s indicated by the similarity of the pygidium to that illustrated by Struve zn Moore (1959) for Trimerocephalus (although the material may not belong to that genus). Both specimens have eleven thoracic segments with a granular prosopon which is also developed on the pygidium. The short, wide and flat pygidia exhibit three axial rings and a terminal piece, and three moderately curved pleural ribs with weak pleural furrows developed laterally. The posterior margin of the pygidium is gently convex, with a narrow marginal roll; the postero-lateral extremities are angular. ACKNOWLEDGEMENTS Thanks are due to Robert Jones (Australian Museum) and John Irving (amateur collector, Merrylands) for making this material available for study, and Dr Noreen Morris (University of Newcastle) for comments on the specimens of Leptophloeum. Research on Devonian faunas has been supported by the ARGS and the University of Wollongong. References BENSON, W. N., 1913. — The geology and petrology of the Great Serpentine Belt of New South Wales. Part 1. Introduction. Proc. Linn. Soc. N.S.W. 38: 490-517. CHESNUT, W. S., FLOOD, R. H., MCKELVEY, B. C., and CAMERON, R. G., 1983. — Manilla 1:250,000 Geo- logical Serves, Sheet SH56-9 (1st Ed.). New South Wales Geological Survey. HAHN, G., and Hann, R., 1975. — Die Trilobiten des Ober-Devon, Karbon und Perm. Jn KROMMELBEIN, K., (ed.), Leztfossilien, begriindet von Georg Gurich. 2nd ed., 1. Berlin-Stuttgart: Gebrider Borntraeger. JENKINS, T. B. H., 1969. — Devonian of the Keepit Inlier. Jn PACKHAM, G. H., (ed.), The geology of New South Wales. J. geol. Soc. Aust. 16: 242-243. ——,, 1972. — Tamworth Trough. Jn ROBERTS, J., e¢ al. (1972): 480-481. Fig. 2. Phacopinid, genus and species indeterminate. A, B, Australian Museum (AM) F65555. A, thorax and pygidium, x2.5; B, pygidium, x6. C, D, AM F72560. G, internal mould, thorax and pygidium, AM F72560b, x2; D, latex cast of external mould of pygidium, AM F72560a, x6.5. Both specimens from Mandowa Mudstone, near Barraba, New South Wales; probably to [V-VI (Late Famennian) in age. PROC. LINN. SOC. N.S.W.,, 110 (1), (1987) 1988 30 LATE DEVONIAN TRILOBITES Mory, A. H., 1982. — A review of Early Carboniferous stratigraphy and correlations in the northern Tamworth Belt, New South Wales. Proc. Linn. Soc. N.S.W. 105: 213-236. PICKETT, J. W., 1960. — A clymeniid from the Wocklumeria Zone of New South Wales. Palaeontology 3: 23a 2A RICHTER, R., and RICHTER, E., 1955. — Oberdevonische Trilobiten Nachtrage. 1. Trilobiten aus der Prolobites-Stufe Il. 2. Phylogenie der oberdevonischen Phacopidae. Senck. leth. 36: 49-72. ROBERTS, J., e¢ al., 1972. — Correlation of the Upper Devonian rocks of Australia. J. geol. Soc. Aust. 18: 467-490. STRUVE, W., 1959. — Suborder Phacopina. Jn MOORE, R. C., (ed.) Treatise on Invertebrate Paleontology Part O. Arthropoda: 0461-495. Lawrence, Kansas: Geological Survey of America and University of Kansas Press. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 Studies in Upside-down Flies (Diptera: Neurochaetidae) Part I. Systematics and Phylogeny DAVID K. MCALPINE McALPINE, D. K. Studies in upside-down flies (Diptera: Neurochaetidae). Part I. Systematics and phylogeny. Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 31-58. Neurocytta and Neurotexis are described as new subgenera of Neurochaeta. Neurochaeta (Neurochaeta) magnifica (Papua New Guinea), Neurochaeta (Neurochaeta) capilo and Neuro- chaeta (Neurochaeta) parviceps n.spp. (West Malaysia) are described. The genus Nothoasteza is transferred from Astelidae to Neurochaetidae and Nothoasteva clausa n.sp. (Western Australia) is described. The ground-plan characters for the Neurochaetidae are given in the light of the newly included taxa, and variation in the structure of the arista, the prosternum, the scutellum, and the postnotum is examined. Phylogeny within the family is discussed. Keys to genera, subgenera, and species are given. David K. McAlpine, The Australian Museum, Box A285 Sydney South, Australia 2000; manuscript received 21 May 1986, accepted for publication 22 July 1987. INTRODUCTION Since the family Neurochaetidae was established for the living Neurochaeta McAlpine and the Baltic amber fossil Anthoclusta Hennig (McAlpine, 1978), more material and information have become available. Field studies of the Australian species have been carried out (Shaw, Cantrell, and Houston, 1982; Shaw and Cantrell, 1983a, 1983b; author’s observations). Woodley (1982) has described two Oriental species of Neurochaeta and made a phylogenetic analysis for the genus. The author’s work (with the help of K. C. Khoo) has brought to light additional species from Malaysia together with information on apparent host plants, and J. W. Ismay has discovered a species in Papua New Guinea. A further specimen of the little known genus Nothoastera Malloch has be- come available, thanks to A. C. Postle and B. Cantrell, and evidence has been obtained indicating that it should be transferred from the Asteiidae to the Neurochaetidae. The number of recognized living species of the family now stands at 10, while there are 2 fossil species described by Hennig (1965, 1969). The morphological terminology here used has been largely outlined by McAlpine (1973a) or is given by Colless and McAlpine (1970). While the identity of the protandrial sternites (those of abdominal segments 6-8) in the Schizophora is now agreed upon by a number of workers (e.g. Crampton, 1942; Griffiths, 1972), the identity of certain other postabdominal structures remains con- troversial. I am quite sceptical about many attempts to homologize structures over a wide spectrum of the order Diptera and even more so with attempts at homology between Diptera and other orders. I use the terms epandrium, hypandrium, surstylus, and gonite to designate structures readily identified by their positions, but because of the great variability they exhibit, I have no confidence that the two latter terms are con- sistently applied to homologous structures, and this applies also to the popular Comstock-Needham nomenclature for wing veins. FAMILY NEUROCHAETIDAE The addition of the genus Nothoasteia makes my previous characterization of the family (McAlpine, 1978) inadequate. I do not herein give a revised characterization PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 By STUDIES IN UPSIDE-DOWN FLIES. I because the morphology of Anthoclusza and Nothoasteza is still incompletely known, but the characters given below for Nothoastera may be taken as an extension of the family characters. Alternatively, a statement of the groundplan characters for Neurochaetidae is difficult, because the precise phylogenetic position of Nothoasteza is unclear, on account of its highly autapomorphic and incompletely known morphology. It is therefore not known to what extent the groundplan characters for Anthoclusia + Neurochaeta (largely typified by Anthoclusza), are also those for Nothoastera + Anthoclusia + Neurochaeta. The following characterization is therefore provisional. Fig. 1. Neurochaeta inversa (based on a photograph from life). Groundplan characters of Neurochaetidae General characters as for superfamily Asteioidea (McAlpine, 1978); postvertical bristles parallel to slightly divergent, posteriorly directed; vibrissa and peristomal bristles present; antennal segment 2 cap-like, with dorsal slit; segment 3 deflexed; PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 33 labella of proboscis broad; postnotal region elevated; with relatively small subscutellum; prosternal plate (basisternum) narrow, joined to angular median prominence of the well defined furcasternum,; the following thoracic bristles present: humeral, 2 notopleural, 2 dorsocentral, supra-alar, postalar, 2 unequal scutellars, 2 upper posterior sterno- pleurals; the following bristles absent: presutural, posterior intra-alar, propleural; hind leg longer than other legs; mid coxae approximated; mid femur significantly shorter than hind femur and more slender than fore femur; costa broken only at end of sub- costa; subcosta indistinct distally; second basal, discal, and anal cells complete; vein 6 sclerotized beyond anal cell but not reaching margin; alula distinct; preabdominal ster- nites broad; tergite 6 of O unreduced; cerci distinct and separate in both sexes. Morphology of the arista I have previously discussed antennal structure in the Asteioidea (McAlpine, 1978; 1983). The arista of Neurochaeta inversa is of a somewhat reduced type: segment 4 is not discernible, segment 5 is very short and annular, and segment 6 has long branches, some of the dorsal ones arising close to its base. This is not in agreement with the arista of Anthoclusia gephyrea as figured by Hennig (1965: fig. 247B), which appears to have a some- what elongate, cylindrical segment 5, and segment 6 with branching less developed at its base. On the whole dry specimen, the arista of Nothoasteva clausa appears to have segment 5 almost as short as in Neurochaeta inversa, but segment 6 has no major branches, only short hairs. A more thorough study of the morphology of the arista in Neurochaetidae will be necessary to evaluate its use in determining phylogeny. Morphology of the prosternum As the prosternal characters are of both taxonomic and adaptational significance, their variation 1s here described. The prosternum of the Neurochaetidae is characterized by reduction, through narrowing, of the basisternum (the sternal plate which is well developed and distinct in most higher Diptera); and the angular anterior production of the furcasternum, to the apex of which the basisternum 1s usually joined. A greater development of prosternal structures is seen in Cyamops (Fig. 3), a genus of the apparently related family Periscelididae. The furcasternum, often invisibly fused with the sternopleura (katepisterna) in other Schizophora, is defined by a V-shaped suture and has a distinct furcal pit in its posterior angle. The very broad basisternum is joined posteriorly to the furcasternum and anterolaterally to precoxal bridges of the propleura. The obvious suture on the precoxal bridge of each side possibly indicates that this bridge is an apomorphic development within the family Periscelididae, as it is absent in Scutops and Persscelis. The periscelidid genus Stenomicra (Fig. 2) has a somewhat similar prosternum to that of Cyamops, but the sutures defining the fucasternum from the sternopleura are not visible in the examples studied, though the cuticle is sufficiently transparent to show any sutures. All these forms have a distinct median line or groove on the sternopleura indicating the internal ridge which extends posteriorly from the furcal pit. Neurochaeta inversa (Fig. 4) retains a large, convex, well defined furcasternum, but the furcal pit is evident only as the angular junction of the grooves along the suture delimiting the furcasternum from the sternopleura. The basisternum is much reduced in size, but not linear, with a marked median groove. In some individuals (more fre- quently in ones from northern populations) there is a fine hair in the sternal membrane on each side of the basisternum. PROG. LINN. SOG. N.S.W., 110 (1), (1987) 1988 34 STUDIES IN UPSIDE-DOWN FLIES. I Figs 2-6. Prosterna of periscelidids and neurochaetids. 2. Stenomicra sp. (Mount Wilson, N.S.W.). 3. Cyamops dayi Khoo. 4. Neurochaeta inversa. 5. Neurochaeta parviceps. 6. Nothoasteva clausa (showing margin of fore coxa at each side). bs = basisternum. fs = furcasternum. In Neurochaeta parviceps (Fig. 5) and closely related species the basisternum is reduced to a narrow-linear sclerite and the hairs in the adjacent membrane are well developed. In Neurochaeta magnifica the basisternum 1s very broad, with several setulae (Fig. 23). In accordance with the evidence that (a) the reduced, narrowed prosternum is the nor- mal condition of all subgenera of Neurochaeta, and (b) N. magnifica is derived from among the more apomorphic types of the subgenus Neurochaeta, 1 regard this condition as a secondary sclerotization which I shall later relate to other peculiar features of the prothoracic region in this species. Nothoasteia clausa shows the most remarkable degree of reduction of prosternal structure (Fig. 6), but this appears to be an extreme development of the condition seen in Neurochaeta. No trace of the basisternum remains. The furcasternum is angular anteriorly as in Neurochaeta, with relatively broad raised margin; posteriorly it 1s delimited by a simple transverse line and is without the sternal pit. There is no median suture extending posteriorly from this transverse suture (hence no internal median ridge), the sternopleura of each side being indistinguishably fused except towards their posterior extremities. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 39 The prosternum of Anthoclusia is not known in detail. Hennig (1969) merely states for A. remotinervis, that the prosternum 1s apparently not connected to the propleuron by a precoxal bridge, and I found it difficult to interpret the prosternal structure during my brief examination of this specimen. Morphology of the scutellum and postnotum The scutellar region and scutellar chaetotaxy of Anthoclusia gephyrea very probably represent the most plesiomorphic conditions among known neurochaetids. This state- ment is made because A. gephyrea has the thorax at an evolutionary stage which precedes marked dorsoventral compression, because the scutellum and postnotal region resemble those of the periscelidid genera Cyamops (Fig. 7) and Perescelis (the family Periscelididae being the apparent sister-group of Neurochaetidae if it is monophyletic), because the general morphology and time level of A. gephyrea indicate that it is a very plesiomorphic neurochaetid; and because the variations in scutellar chaetotaxy in the genus Neurochaeta are readily explained as simple or serial derivatives from it. The scutellum of A. gephyrea (Hennig, 1965: fig. 248) is subtriangular with a broadly rounded apex; it appears to be somewhat flattened on the dorsal surface and quite thick at the free margins. There is a small lateral pair of bristles situated some dis- tance behind the scutellar suture, and a large subapical pair of bristles. Because of varia- tion in the position of the latter pair, they are hereafter referred to as the major pair of scutellar bristles. It is uncertain if their suberect position shown in Hennig’s figure 1s natural or not. The scutellum of Anthoclusia remotinervis differs from the above in loss of the lateral pair of bristles (Hennig, 1969). In the genus Neurochaeta the scutellum has become markedly shortened and more or less depressed relative to that of Anthoclusia gephyrea, but otherwise the structure of the scutellum exhibits diverse levels of specialization among the species. In Neurochaeta prisca (McAlpine, 1978: fig. 11) the outline of the scutellum retains something of the plesiomorphic triangular shape despite the shortening, and the major bristles remain close together near the apex. Each lateral bristle is replaced by a series of setulae, an apomorphy unique in the family. The remainder of the species of Neurochaeta (subgenera Neurotexis and Neurochaeta) have the scutellum usually with evenly rounded outline (tending towards a semicircle) and the major bristles are markedly laterally displaced. Neurochaeta (Neurotexis) stucken- berg has retained the scutellum at this level of specialization (McAlpine, 1978: fig. 12). The subgenus Neurochaeta is characterized by acquisition of additional bristles or setulae between the major pair, on or near the posterior margin. These I term the posterior scutellar bristles for descriptive purposes. The tendency towards a more erect position of the bristles is apparent throughout the subgenus, but the variation, with retention of symmetry, seen in dried material of Neurochaeta capilo and, especially, N. inversa seems to indicate much mobility of the major pair in life. In N. capilo (Fig. 10) there is one pair of short convergent bristles near the apex of the scutellum and few, often asymmetrically placed setulae near them. This somewhat irregular arrangement may represent an approximation to an early stage of evolution of these posterior macrotrichia. In N. inversa (McAlpine, 1978: fig. 16) the posterior macrotrichia are represented by a regular linear series of rather short bristles along the posterior margin of the scutel- lum, in two to four (usually three) pairs. In N. magnifica the arrangement of the posterior bristles is similar, but there are four to seven pairs. The scutellum in the parviceps group (Figs 9, 11) has the major bristles shorter and more consistently erect (or less posteriorly inclined) than those of N. capzlo and N. inversa, PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 36 STUDIES IN UPSIDE-DOWN FLIES. I 12 Figs 7-12. Scutella and postnota of periscelidids and neurochaetids. 7. Cyamops dayt (posterior aspect). 8. Neurochaeta capilo (posterior). 9. Neurochaeta macalpinei (posterior). 10. Neurochaeta capilo (dorsal). 11. Neurochaeta parviceps (dorsal). 12. Nothoasteia clausa (dorsal). su = subscutellum. In Fig. 11 bristles shown as if extended horizontally. and the posterior bristles represented by one pair, each of which is closer to the major bristle than to the other posterior bristle. This arrangement of the posterior bristles may have been evolved through condensation of the linear series seen in N. inversa, or, PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 D. K. MCALPINE Sr, alternatively, it may have been attained directly from the less ordered configuration of N. capilo. In the latter case, N. inversa and the parviceps group have acquired independent apomorphic conditions of the posterior bristles. Newrochaeta parviceps has all three pairs of scutellar bristles more similar in length than in other species of the group. (In Fig. 11 the suberect bristles of the major pair have been drawn as if in a horizontal plane.) N. sabroskyi probably has the scutellar bristles similar to those of N. parviceps, but in the unique holotype only the lateral bristles are preserved. N. macalpine: has many of the body bristles shortened as an autapomorphy, but it is not clear precisely how this general process has affected the scutellar bristles. In contrast to N. parviceps, the major bristle in N. macalpinei (Fig. 9) is about twice as long as the lateral bristle or the posterior bristle, but the major bristle is only slightly shorter than that of N. parviceps relative to the dis- tance between the bases of the major bristles. This size differentiation may be the re- tention of a plesiomorphic condition, as perhaps 1s that of N. capilo, or a late apomorphy derived from the condition seen in N. parviceps, out-group comparison (for the parviceps group) suggesting the former. In N. macalpine: the major bristles are strongly convergent and their backward inclination is minimal. The scutellum of Nothoasteia clausa (Fig. 12) has some points of resemblance to that of the parviceps group of Neurochaeta; in particular it is flat, with rounded posterior margin but 1s even shorter; there are 3 pairs of bristles though these are all much shorter than in the parviceps group and directed posteromedially. On its appearance, without reference to other facts, the scutellum might be considered to be derived from that of the parviceps group, but, as shown below this is negated by other evidence. The postnotal structure of the periscelidid genus Cyamops (Fig. 7) is taken as representing a generalized condition near that of the groundplan of the Periscelididae- Neurochaetidae (and perhaps of the Asteioidea), which preceded dorsoventral com- pression. Immediately below the free posterior margin of the scutellum there 1s a narrow transverse zone of membranous cuticle, the subscutellar membrane. The median sclero- tized part of the postnotum (mediotergite), lying below the scutellum, consists of two distinct convex zones, the smaller upper subscutellum and the lower postscutellum, which slopes posteriorly to the first abdominal tergite. Postnotal structure in Anthoclusia gephyrea appears not to differ from the above condition so far as I can discern from the imperfect detail in the illustration by Hennig (1965: fig. 248). In particular the posterior margin of the scutellum remains quite deep and the subscutellum is markedly smaller than the well developed postscutellum. Other neurochaetids all have the postnotal region reduced in depth, apparently as a result of dorsoventral compression of the thorax. Unfortunately I no longer have study material of Neurochaeta prisca for detailed comparison. My previously recorded observations (McAlpine, 1978: 282) indicate a sig- nificant reduction in depth of the postnotum without its acquiring a prominently convex condition. This perhaps indicates that much of its remaining surface consists of post- scutellum rather than subscutellum, in contrast to the other species of the genus Neurochaeta. Interpretation of postnotal structure in the subgenus Neurochaeta is aided by refer- ence to its most plesiomorphic species, N. capilo (Fig. 8), but the structure of the latter has evolved far beyond the apparent neurochaetid groundplan and no known neurochaetid represents a clearly intermediate stage. The morphologically diverse periscelidid genus Stenomicra shows a range of post- notal structure which parallels some of the probable evolutionary stages of neuro- chaetids between the groundplan condition and that of N. capilo. In Stenomicra sp. (West Malaysia, Australian Museum, a remarkably plesiomorphic representative of the genus), the condition is rather similar to that of Cyamops as described above, but the PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 38 STUDIES IN UPSIDE-DOWN FLIES. I subscutellum has become enlarged at the expense of the postscutellum. In other species of Stenomicra (subgenera Podocera and Stenomicra) the subscutellum becomes very convexly prominent and increases further in depth. It approaches the first abdominal tergite on the median line in extreme cases, thus almost dividing the postscutellum in two. Neurochaeta capilo has the postnotal region (Fig. 8) somewhat resembling the condition described for the more apomorphic species of Stenomicra, but there has been a decrease in depth and the subscutellum is broader, almost as wide as the postscutellum, strongly convex, with narrowly acute lateral extremities. The visible demarcation between subscutellum and postcutellum is not sharp, partly from the dense clothing of silvery pruinescence, but it appears that the subscutellum is only narrowly separated from the first abdominal tergite on the median line by the medially narrowed postscutel- lum. The upper margin of the subscutellum, where it meets the subscutellar membrane, is incurved in conformity with the general convexity of the surface, but is not abruptly inflexed (Fig. 13). ia 15 16 Figs 13-16. Diagrams of scutellar and postnotal structures in Neurochaeta in vertical longitudinal section, scutellum at top, postnotum at left, subscutellar membrane indicated by thin line. 13. N. capilo. 14. N. inversa. 15. N. magnifica. 16. N. parviceps. The inversa, magnifica and parviceps groups of Neurochaeta have the subscutellum both deeper and broader than in N. capilo and the postscutellum thus reduced to two small lateral pieces (Fig. 9). In the inversa and magnifica groups, but not in the parviceps group, the upper margin of the subscutellum is abruptly inflexed, so that there is a trough between the scutellum and subscutellum, the floor of which is formed by the subscutellar membrane (Figs 14, 15). I no longer have access to material of Neurochaeta stuckenbergi. Clearly this species has a strongly convex postnotal region largely developed from the postscutellum but other details were not recorded. Again there is resemblance in postnotal structure between Nothoasteia and the most apomorphic species of Neurochaeta. Nothoastera clausa has the subscutellum very convex and shallow, but almost entirely filling the area between the scutellum and the base of the abdomen, the postscutellum being reduced to two small lateral plates. No subscutel- lar membrane is visible in the only specimen, the sclerotized subscutellum being appressed to the ventral part of the scutellum. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 39 Hardy (1950) has recorded what he termed ‘the articulating scutellum’ in a number of brachycerous Diptera, including several of the Schizophora, where the condition is apparently of wide occurrence. Examination of series of dried specimens of Neurochaeta inversa and N. parviceps indicates that movement occurs between the scutellum and sub- scutellum. The subscutellar membrane becomes infolded when these parts are appressed, and the margin of the scutellum becomes partly enveloped by the upper margin of the subscutellum. In the available specimen of Nothoasteva clausa the upper margin of the subscutellum passes beneath the scutellum, but does not envelop it. The relations of these parts are shown diagrammatically in Figs 13-16. Phylogeny The apparent phylogenetic lines in the family, leaving out of consideration Nothoasteia and the new species of Neurochaeta here described, are given by Woodley (1982). It remains for these taxa to be added to Woodley’s system. At the time of Woodley’s work it appeared that the apomorphic condition of charac- ter 3 (posterior cubital or anal cell reduced or absent) was a groundplan state for Newro- chaeta stuckenbergx + the N. inversa group (1.e. subgenera Neurotexts and Neurochaeta in the present work). The newly discovered species N. capzlo is undoubtedly a member of the latter group of this pair but has a complete anal cell and sclerotized vein 6 beyond it. It seems, then, that reduction of the anal cell (and vein 6) has taken place independently in these two subgenera. I believe evidence for sister-group relationship between subgenera Neurotexis and Neurochaeta is confirmed by the following apomorphic characters in re- lation to the subgenus Neurocytta (including N. prisca): costa with subcostal break obliquely incised to produce a lobe; subscutellum deep and convexly prominent; major (primarily apical) pair of scutellar bristles displaced laterally; only one upper posterior sternopleural bristle present. The revised set of distinctive apomorphic characters for the subgenus Neurochaeta (the ‘NV. inversa species group’ of Woodley, expanded to include new species) is as follows: fronto-orbital bristles reduced to 3 pairs; postvertical bristles lost; a series of postgenal bristles present; suborbital bristle present (below lowest point of eye, see Figs 18, 26); second (from rear) dorsocentral bristle reduced and approximated to prescutellar dorso- central, or lost; scutellum with at least one pair of posterior marginal bristles or hairs between major pair; sternopleuron anteriorly with group of strong hairs or bristles; macrotrichia on radial sector developed as about 3 strong bristles; second basal cell confluent with first basal cell; ©: surstylus detached from margin of protandrium. The subgenus Neurochaeta includes the capzilo, inversa, magnifica, and parviceps groups, as newly defined here. The position of Neurochaeta capilo as a sister group to the remainder of subgenus Neurochaeta (inversa group sensu Woodley) is evidenced by its having the following plesio- morphic characters relative to those uniformly present in the rest of the subgenus: thorax not strongly depressed; 2 dorsocentral bristles present; anterior sternopleural bristles not differentiated; mid coxae almost contiguous; anal cell closed by distinct, curved anal crossvein; vein 6 developed beyond anal cell; sternite 8 of male large; cercus of Q small, not plate-like. For these reasons I place N. capilo as the only known species of the capilo group. The apomorphic characters differentiating the parviceps group (N. sabroskyi, N. parviceps, and N. macalpinet) are: suborbital bristle curved downwards; prelabrum much reduced; palpus very short; propleuron without distinct callus; prosternum narrow- linear; metasternum extensively setulose; mid coxae separated by at least width of each coxa; mid femur with ventral comb of weak bristles; vein 5 not extending beyond discal cell. The monotypic inversa group lacks the above characters, but has apparently few PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 40 STUDIES IN UPSIDE-DOWN FLIES. I “ (a0) qo o | (a0 oO a0) oO n ‘d iS xe) O Ss Ou = Q, Ss ‘d © ‘A eb) SS oy a0} o e) G4 n a. oO ”Y Cac} O oO Se oH ‘dl iS oH Cal O n > n oO ‘Hl = oO 0} > se Ss} i) ‘d S} OQ. 80 => O iu a 140) (a0 w £5) oO oO cS © © o eS Q, n oO = al = OQ. n O Q. o o oO o oO o o fe) a = a a = a = = = Anthoclusia Oligocene Spp. Eocene Fig. 17. Dendrogram showing apparent interrelationships of neurochaetid taxa. apomorphic characters not shared with the parviceps group, viz.: posterior scutellar bristles forming a regular transverse series; dorsal margin of subscutellum inflexed; ster- nite 8 of O& absent; and (probably) propleural callus enlarged. The group is, however, intermediate between the capilo group and the parviceps group in several characters, viz. degree of development of anterior sternopleural bristles; incipient setulosity of meta- sternum; degree of separation of mid coxae; length of tarsal claws; position of anterior (incurved) fronto-orbital bristle. The magnifica group (also monotypic) has an extraordinary array of apomorphic characters, including all those given for the znversa group except loss of © sternite 8, some of those given for the parviceps group (viz. metasternum extensively setulose; mid coxae separated by at least width of each coxa), and numerous apparently auta- pomorphic characters (e.g. head depressed; inner vertical bristle absent; anterior notopleural bristle displaced dorsally; hypopleuron setulose; prosternum very broad; hind femur much longer than thorax; costal armature simplified). The apparent conflict in phylogenetic evidence from variously shared apomorphic characters and the phenetic remoteness from other species of subgenus Neurochaeta complicate the problem of the immediate relationships of N. magnifica. Careful consideration leads me to the conclu- sion that those characters shared with the inversa group are most likely to be true synapo- morphies (although the first two are present in an exaggerated form in N. magnifica), while the characters shared with the parviceps group are convergent. The broadly ex- posed, setulose metasternum and widely separated mid coxae are associated with a PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 41 broadening and flattening of the thorax in a number of separately derived dipterous types (e.g. Coelopa, Orygma, the pupiparous flies). That a broadly flattened thorax has been achieved to some extent independently in the parviceps and magnifica groups is evidenced particularly in the position of the fore coxae and their relation to the pleura. In the face of contrary evidence, then, the shared apomorphic characters in the two groups do not carry conviction as true synapomorphies. I therefore postulate a sister- group relationship between the inversa and magnifica groups. This is in accordance with geographic distribution, as these are the only groups of Neurochaeta in the Australian (Australasian) Region. The phylogenetic position of the genus Nothoasteia in relation to other neurochaetids is difficult to determine. The genus was originally placed in the family Asteiidae by Malloch, 1936, with considerable doubt. The following is a list of those characters for Nothoasteia, which have influenced me in transferring the genus from the Astelidae to the Neurochaetidae. 1. Antennal segment 2 cap-like, with dorsal slit (not cap like, with dorsal part of distal margin slightly sinuate only, in Asteudae). 2. Upper occiput without trace of deep, broad concavity characteristic of Asteiidae (including Succznastera and Leiomyza). 3. Prothoracic basisternum greatly reduced in a broad, membranous intercoxal field; furcasternum with acute anterior lobe. These characters are peculiar to the Neurochaetidae among the Asteioidea (see further discussion above). The Asteiidae have the prosternum broad, sometimes little sclerotized, joined to the almost straight, transverse anterior margin of apparent mesosternum. 4. Metasternum separated from hypopleuron on each side by membrane of hind coxal cavity (narrowly joined to hypopleuron in Neurochaeta, usually very broadly joined in Asteidae). 5. Fore coxa short (generally relatively long and slender in Asteiidae). 6. Mid coxae very short, separated (mid coxae contiguous and relatively prominent in Asteudae). 7. Hind coxal cavities directed posteriorly to posterolaterally (directed ventrally in Asteiidae). 8. Hind legs (particularly hind femora) enlarged; mid legs reduced in size. This is a slight extension of the highly characteristic leg proportions seen in other neuro- chaetids. (Asteiidae never have such disparity in leg size.) 9. Vein 6 more complete than in any known asteiid (including Succinastera). 10. Abdominal sternites broader than in any asteiid known to me (including Succinastera), despite the more slender abdomen. The broadened sternites are particu- larly characteristic of the Neurochaetidae. In making the above comparison I have referred to Hennig’s (1969) detailed description of the Baltic amber fossil Succinasteia carpenter, which could represent the stem group from which all recent Asteiidae were derived. I have also examined Bryania bipunctata Aldrich and an undetermined asteiid from Papua New Guinea, both of which have, like Nothoasteia, a somewhat depressed body form. They do not, however, show significant resemblance to Nothoastza in other characters. Nothoasteva resembles the Asteiidae (or at least some of its species) in its small size, reduced chaetotaxy, and reduction in wing venation, all characters where convergence is most apt to occur, the two latter states often accompanying size reduction in various acalyptrate groups. The absence of a costal break, shared by Nothoasteia and Asteiidae, has been considered a plesiomorphic character by Hennig (1958), but, as Griffiths (1972) has pointed out, it is likely to be an apomorphic acquisition in the groundplan of the Asteiidae. It appears to represent an independent autapomorphy in Nothoasteza. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 42 STUDIES IN UPSIDE-DOWN FLIES. I It has been suggested, without reference to specimens, that Nothoasteia may belong in the family Anthomyzidae, but I do not consider that this is a reasonable theory. I know of no significant apomorphies shared between it and the anthomyzids, and Nothoasteia differs in characters of antenna, prosternum, leg proportions, and mid coxae, much as it differs from Astelidae. Possibly, Anthomyzidae and Astelidae are more closely related to each other than either 1s to Nothoastera and the Neurochaetidae. Within the Neurochaetidae, Nothoasteia resembles Anthoclusia more than other known forms in the development of antennal segment 2. This would appear to exclude it from immediate relationship with subgenera Neurotexis and Neurochaeta which share the enlarged, hood-like segment 2, probably as a synapomorphy. Unfortunately the antenna of subgenus Neurocytta, evidently a sister group to the rest of genus Neurochaeta, is unknown. The probably plesiomorphic approximation of the apical scutellar bristles in Nothoasteia seems to confirm its exclusion from this Neurotexis-Neurochaeta complex. Possibly the more cylindrical abdomen, excludes Nothoasteza from close relationship with the whole genus Neurochaeta, but this character needs further study. The confluence of the second basal and discal cells in both Nothoastera and subgenus Neurotexis appears to be due to convergence. This is a condition which has been evolved many times in the acalyptrates. It does, however, confirm the distinction of Nothoasteia from subgenus Neurochaeta, in which the second basal cell is consistently separated from the discal cell and confluent with the first basal cell. Nothoasteia resembles species of subgenus Neurochaeta in several characters which are associated with flattening of the thorax. The partly almost planate sternopleuron with a series of upper, anterior bristles is such a character, as is the short, flat scutellum and bulging subscutellum. But in the groundplan of the genus Neurochaeta the thoracic flat- tening is less developed, as are these associated apomorphic characters. This indicates a complex convergence between Nothoastera and the more apomorphic forms of Neuro- chaeta. Loss of certain bristles, e.g. the postvertical and supra-alar, also constitutes con- vergence between Nothoasteia and subgenus Neurochaeta, but, on the whole, reduction in chaetotaxy has progressed further in the former. I conclude from the above morphological evidence that Nothoasteia separated from the main neurochaetid stem certainly before the divergence of subgenera Neurotexis and Neurochaeta, probably before the separation of subgenus Neurocytta, and possibly earlier than the stage reached by the fossil Anthoclusia. The very marked autapomorphies of Nothoasteia (including the flattened head, setulose eyes, presence of mesopleural setulae, modifications of the furcasternum, sternopleura, and metasternum, loss of tarsal claws, loss of costal break, almost total fusion of subcosta and vein 1, loss of alular fringe, divided sternite 7 of the female abdomen) suggest that the separation from other forms may be particularly early in the history of the Neurochaetidae. Those apomorphies have, however, had a transforming effect on the insect as a whole and make its accurate placement in the phylogenetic system impossible at present. Geographic origins The Neurochaetidae are at present distributed in wetter parts of the Old World tropics and subtropics, and there is no evidence that they have occurred in the New World. Anthoclusia, which fulfils morphological requirements for a stem group to genus Neurochaeta (and possibly to Nothoasteia), occurred in northern Europe in the late Eocene or the commencement of the Oligocene. The genus Neurochaeta was apparently derived from a species which had acquired further apomorphies (McAlpine, 1978). Anthoclusia lived at a time when Africa was still separated from Europe by the Tethys Sea, and judg- ing from the meagre but probably significant evidence from mammalian fossils PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 43 (Coryndon and Savage, 1973), European land fauna did not invade Africa until well into the Oligocene. If early neurochaetids were strictly inhabitants of moist forest, like their descendants, they were probably no more vagile than land mammals. It is probable therefore that Anthoclusia was restricted to Eurasia at the Eocene-Oligocene boundary, and the two separate lineages (subgenera Neurocytta and Neurotexis), which are now in the Afrotropical Region, reached there some time later in the Tertiary. Until better collec- tions of Afrotropical neurochaetids are made no significant comparison can be made between the faunas of Madagascar and the African mainland. The restriction of the Australasian species to the vicinity of rainforest in tropical and subtropical areas, together with their similarity to oriental species, indicates a late arrival in Australasia, perhaps during the Pliocene when sea barriers to the north were at aminimum. The distribution of Nothoasteza is little known, but its occurrence in southwestern Australia suggests that the genus 1s part of an older Australian fauna, without indicating more precisely the time of its occupation. KEY TO RECENT GENERA OF NEUROCHAETIDAE Eye setulose; costa unbroken; alula without fringe; functional tarsal Gla Seal lo) S © In ten tere eal eet MU ea pel AEs MR re ee Meret Rise gi Nothoasteia Eye without obvious setulae; costa with break at end of subcosta; alula with marginal fringe; tarsal claws developed .................. Neurochaeta GENUS NEUROCHAETA MCALPINE Neurochaeta McAlpine, 1978: 278-281. Type-species N. inversa McAlpine. Five species have been added to this genus since its description (Woodley, 1982, and the present work), but these are all rather closely related to the type-species and are referable to the same subgenus. Despite doubt as to the phylogenetic position of the species of Nothoasteza in relation to the Neurochaeta species, there is no persuasive evidence that Neurochaeta s.l. is polyphyletic. The afrotropical species, N. prisca and N. stuckenbergz, are at present known to me from a miniscule sample of three specimens, which may not be sufficient to indicate the diversity of the fauna and consistency of group characters. Nevertheless, as shown above, these species are phylogenetically isolated from other known neurochaetids and are now considered to require their own subgenera. Key to subgenera of Neurochaeta 1 Second basal cell completely enclosed; upper posterior sternopleural bristles 2; scutellar bristles of major pair approximated near apex of Seice! MMMeESMIOS CUCE LUI) NOt PLOUMIIEIG 4) sere eaere cope Neurocytta — Second basal cell incomplete; upper posterior sternopleural bristle solitary; scutellar bristles of major pair widely separated; subscutel- OVEN KEI? COMIS:< ZUaXGl JONAOVETUNS TE Je Soa abonounchsanapecogea nes 2 2 Second basal cell confluent with discal cell; costa terminating at vein 3; radial sector without bristles; postvertical, presutural, and supra- alar bristles present; suborbital bristle absent; posterior margin of Scuitell umm aren etweehuia) Or bGistless erg eee. es sees Neurotexts — Second basal cell confluent with first basal cell; costa terminating at vein 4; radial sector towards base with 2 to 4 strong erect bristles; presutural, supra-alar, and usually, postvertical bristles absent; suborbital bristle usually present; posterior margin of scutellum with one or more pairs of shorter bristles between major pair ..... Neurochaeta PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 44 STUDIES IN UPSIDE-DOWN FLIES. I Subgenus Neurocytta n. subg. Type species: Neurochaeta prisca McAIpine. Fronto-orbital bristles 4, one of which is inserted mesad of and a little behind fore- most of the aligned series; short postvertical bristle present; cheek with one postgenal bristle situated behind lowermost point of eye (not homologous with suborbital bristle of subgenus Neurochaeta). Presutural bristle absent; one dorsocentral bristle present; supra- alar bristle present; scutellar bristles of major pair approximated near apex of scutel- lum; scutellum otherwise with short lateral hairs only; postnotum receding below scutel- lum, not prominent; sternopleuron with 2 upper posterior bristles and no anterior bristles. Wing with strong, spaced anterior bristles on costa between terminations of veins 1 and 2; costa extending to termination of vein 4, with simple break at termination of subcosta; macrotrichia on radial sector represented by small hairs on stem of veins 2 and 3 dorsally; second basal cell separated from both first basal and discal cells; anal cell completely enclosed; vein 6 well developed beyond anal cell; alula relatively broad. Postabdomen of male with large, asymmetrically placed sternites 6 and 7, dorsal sternite 8, and well developed epandrium; cercus in both sexes inconspicuous, not plate-like. The distinctive characters of the only included species of Neurocytta, viz. Neurochaeta prisca, and their phylogenetic significance have been discussed by McAlpine (1978) and Woodley (1982). This species is still only known from the type material from Zimbabwe. The subgeneric name is derived from the Greek vevgov, nerve or wing-vein, and xUTTAQOS, cell in reference to the well developed cells in the basal part of the wing. It is feminine on account of the ending. Subgenus Neurotexis n. subg. Type species: Neurochaeta stuckenberg: McAlpine. Fronto-orbital bristles 4, the inclinate one inserted mesad of and at same level as anterior reclinate one; postvertical bristle present; one postgenal bristle as in Neurocytta; suborbital bristle absent. Presutural bristle present; dorsocentral bristles 2; supra-alar bristle present; scutellar bristles in 2 pairs: widely separated major pair, and shorter lateral pair; subscutellum deep and convexly prominent; sternopleuron with one upper posterior and no anterior bristles. Costa terminating immediately beyond end of vein 3, with fairly strong spaced anteroventral bristles between terminations of veins 1 and 2, without short erect dorsal setulae immediately before end of subcosta; costal section be- fore subcostal break produced distally in front of break as a long, finger-like process; radial sector without macrotrichia; second basal cell separate from first basal cell, con- fluent with discal cell; anal cell small, imperfectly enclosed; vein 6 little developed beyond anal cell; alula rather narrow. Postabdomen of male unknown; cercus of female narrow, subcylindrical. The characters and relationships of the only included species are discussed by McAlpine (1978) and Woodley (1982). It is only known from the unique holotype from Madagascar. The subgeneric name is derived from the Greek vevgov, a nerve or wing-vein, and Te&.s, a wasting or dissolution, in reference to the distal fading of the costa and vein 4. It is feminine. Subgenus Neurochaeta s. str. Fronto-orbital bristles 3, the inclinate one situated in front and mesad of foremost reclinate one; postvertical bristle absent (except in N. magnifica); a variable series of post- genal bristles present along posterior margin of eye; suborbital bristle present below lowest point of eye (Figs 18, 26; reduced in N. magnifica, Fig. 25); one dorsocentral or 2 PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 45 approximated bristles present; supra-alar (as distinct from postalar) bristle absent; scutellar bristles of major pair sublateral on posterior margin, with shorter intermediate bristles or hairs on posterior margin and pair of anterolateral bristles; postnotum largely occupied by the convexly prominent subscutellum; sternopleuron with one upper posterior bristle and with either a variable series of bristles or a number of bristly hairs anteriorly. Wing with variable, usually rather weak, spaced anterodorsal and antero- ventral bristles on costa between terminations of veins | and 3; costa extending to end of vein 4, often with obliquely incised break at termination of subcosta, but not with long process as in Neurotexis; macrotrichia on radial sector represented by normally 2 or 3 strong bristles; second basal cell confluent with first basal cell, separated from discal cell; anal cell and vein 6 variably developed, the former always narrow; alula moderately developed or rather narrow. Asymmetrical sternites of protandrium reduced or absent; epandrium reduced to a transverse, usually medially narrowed plate; surstylus detached from margin of protandrium. This subgenus includes 6 known species from the Oriental and Australian Regions. Key to species of subgenus Neurochaeta 1 Head depressed, longer than high; inner vertical bristle absent; anterior crossvein well before middle of length of discal cell; prosternum very broad; hypopleuron setulose; hind femur much longer than thorax; host apparently Pandanus (magnifica group) ... . magnifica — Head not longer than high; inner vertical bristle well developed; anterior crossvein near or beyond middle of length of discal cell; prosternum attenuated; hypopleuron bare; hind femur not longer (Caveat HOVON cease dl GVRIRe ae eee nett ach anna amy eT ae ee ee laa Deni CN cen 2 2 Vein 5 not continued beyond discal cell; anterior sternopleural bristles in a well developed series, pale yellowish; prosternum very narrowly linear; metasternum extensively setulose; mid coxae separated from one another by at least the width of each coxa; mid femur with loose comb of ventral bristles on basal half; scutellum with one pair of shorter bristles between major pair of bristles on posterior margin; suborbital bristle curved downwards; ©’ postabdomen with sternite 8 present but very short (condition unconfirmed for N. SAUDOSSIG)) ((VAOHAZES (HOD) 4s olin oes eo 8 Ob oso dense oma aoe ee 3 — Vein 5 continued beyond discal cell; spaced dorsal setulae of costa fine and sharp; anterior sternoplural bristles black or undifferentiated; prosternum lanceolate; metasternum bare or with minute incon- spicuous setulae; mid coxae separated from one another by much less than width of each; mid femur without distinct ventral comb; scutellum with 2 or more pairs of bristles or setulae between major pair of bristles on posterior margin; suborbital bristle curved outwards; ©’ postabdomen with sternite 8 nearly as long as tergite 6 OVC BION STAE ease enti ana) a RnB cas Ses Can een Suma ea Ie ee 5 3 Femora pale yellowish, except for dark apical part of hind one; fore coxa not distinctly keeled; abdomen with large pale yellowish zone covering tergites 1, 2, and anterior margin of 3; length of dorso- central bristle not more than half length of scutellum; discal cell at distal end slightly narrower than marginal cell at same level; host BONA NIP AUNTIE civic 4 diotdane ad 6 BD, Oh BIN Big ae Oe among NIE macalpinet PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 46 STUDIES IN UPSIDE-DOWN FLIES. I — Femora largely black; fore coxa with strong anterodorsal keel distally; abdomen with pale yellowish basal zone less extensive or absent; length of dorsocentral bristle more than half length of scutellum; discal cell at distal end slightly broader than marginal cell at same Level ete Sai tRNA eae a luli gas ARAL EKER Vrcid se ML c at peas teva 4 4 Abdominal tergite 1 pale yellowish, contrasting with darker rest of abdomen; capitellum of haltere creamy white; thorax much wider than head; wing less than twice as long as thorax, and less than 3 times as long as wide; host apparently Zingiber................. parviceps — Abdominal tergite 1 brownish, not strongly contrasting with rest of abdomen; capitellum of haltere tawny; thorax about as wide as head; wing more than twice as long as thorax, and more than 3 tumesrasiloncas;wides host umkmowniy aise) er oe ese eee een sabroskyt 5 Sclerotized section of vein 6 present beyond anal cell; anal cell completely enclosed; mid femur without anteroventral bristle; anterior sternopleural bristles not differentiated from hairs; a short second dorsocentral bristle close in front of major one; cercus not broad and plate-like; sternite 8 of O large, nearly as long as tergite Oxshostappanentlye Musa (capelo Croup) renee ee eee capilo — Vein 6 scarcely discernible beyond anal cell; anal cell vestigial; mid femur with short anteroventral bristle; some anterior sternopleural bristles fairly long; only one dorsocentral bristle present; cercus in both sexes broad, plate-like, exserted; sternite 8 of © absent; host Allocastas((nversaicroulp) ise) ee et ee ee eects ae ee inversa Capilo Group n. gr. Suborbital bristle curved outwards; prelabrum moderately developed; humeral callus compact; propleural callus rather small, convex, not sharply defined below; palpus rather long; prementum narrow; thorax not much depressed; scutellum with minor bristles and hairs between major pair of bristles not in a transverse row; upper margin of subscutellum incurved, not inflexed; anterior sternopleural bristles not differentiated from coarse hairs; prosternum rather broadly lanceolate; metasternum bare; mid coxae broad, approximated; mid femur with neither anteroventral bristle nor ventral comb; tarsal claws long and attenuated, each as long as last tarsal segment; vein 5 extending beyond discal cell; anal cell complete; vein 6 developed beyond anal cell; sternite 8 of male large; cercus in both sexes neither plate-like nor posteriorly exserted. This group is at present monotypic. It occurs in the Oriental Region. Neurochaeta capilo n.sp. (Figs 8, 10, 13, 18-21) o Q. Resembling N. inversa and agreeing with description given for that species (McAlpine, 1978) except as indicated below. Coloration: Head greyish brown; parafacial and anterior margin of postfrons orange-fulvous; postocular zone pale grey; antenna and many of its bristles and setulae creamy white; setulae of antennal segment 1 and those on inner surface of segment 2 black; one major dorsal and ventral bristle on segment 2 black or brown; arista beyond base brown. Thorax greyish brown, paler in parts. Hind coxa pale yellowish; other coxae greyish brown; femora brown, fore one pale yellowish basally and apically, mid and hind ones pale yellowish basally; fore tibia yellowish apically; fore tarsus yellowish; other tarsi predominantly brown or hind one paler basally. Wing with whitish zone PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 47 Figs 18-20. Neurochaeta capilo. 18. Head. 19. Wing (ventral). 20. Dorsal sclerites of male postabdomen (not compressed). ep = epandrium. so = suborbital bristle. s8 = sternite 8. t6 = tergite 6. extending to extreme base, but distally extending only slightly beyond forks of veins 2 and 3, and 4 and 5; alula tinged with brown. Haltere pale creamy, more yellowish on scabellum. Head. Height of cheek about 0.15 of height of eye; postfrons a little concave medially but without narrow median channel; eye less oblique than in other species of subgenus; incurved fronto-orbital bristle inserted only slightly anteriorly to anterior reclinate fronto-orbital. Thorax less depressed than in other species of subgenus; 2 dorsocentral bristles present, anterior one short and close to posterior one, sometimes little differentiated; scutellum with bristles of major pair sublateral, also a much shorter lateral and con- vergent posterior pair present in addition to several fine dorsal setulae towards apex; upper anterior sternopleural bristles not differentiated from coarse black hairs of this region. Fore coxa not markedly compressed, without anterodorsal keel; mid femur without anteroventral bristle; hind femur with one long dorsal and one anteroventral (rather than anterior) bristle beyond middle. Costa with black bristle terminating basal section and series of anterodorsal setulae preceding it much stouter than in other species of subgenus, and with pale subterminal dorsal setula absent (or represented by a weak more basally placed setula); costal index 4.8-4.9 (0), 5.1-6.3 (9 ); vein 4 index 2.6-3.0 (o”), 1.9-2.4 (9); anal cell narrow, enclosed distally by curved crossvein; vein 6 well sclerotized for some distance beyond anal cell. PROG. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 48 STUDIES IN UPSIDE-DOWN FLIES. I Abdomen more slender and slightly less depressed than in N. parviceps, tergites 2-6 each with submarginal lateral groove and a bristle on lateral margin which is no more than half as long as tergite. ©’ postabdomen: sternite 6 much reduced, on left side, mirrored by a much smaller sclerite on right side; sternite 7 less reduced than sternite 6, fused to left lateral margin of sternite 8; sternite 8 a little shorter than tergite 6, with several fine setulae; epandrium much narrowed medially, produced at anterolateral angles, with 6 fine setulae on posterior margin; surstylus becoming slender beyond basal plate, with minute apical hairs; outer gonite with posteriorly directed apex and one setula on inner surface; inner gonite much expanded distally into an anterior and a posterior process, with 2 setulae on outer surface; basiphallus short, sclerotized; dis- tiphallus exceedingly long and slender, membranous, with one longitudinal pigmented strip, membranous part towards base with numerous weak, minute denticles, apical part not thickened; cercus elongate, passing forwards from its base and decurved apically, with a series of long bristles along outer surface and fascicle of finer bristles at apex. Q postabdomen: cercus very short and inconspicuous, apparently ovoid, with few, minute setulae. Dimensions: total length, & 2.7-3.2mm, Q 2.6-3.1 mm; length of thorax, o 1.1-1.3 mm, Q 1.0-1.1mm; length of wing, © 2.5-2.8mm, Q 2.3-2.7mm; length of distiphallus c. 3.6mm. Distribution: West Malaysia — Selangor. Holotype 9: Old Bentong Pass, E of Gombak, 15.iv.1985 (Australian Museum), K. C. Khoo and D. K. McAlpine. Faratypes: same locality, 15-16.iv.1985, 14.viii.1986 (1 9 British Museum (Natural History), 2 0,29, Australian Museum), same collectors. Figs 21, 22. Genital segment, ventral aspect (only base of distiphallus shown, setation omitted from right cercus). 21. Neurochaeta capilo. 22. N. parviceps. aa = aedeagal apodeme. c = cercus. dp = distiphallus. eb = epandrial bristle. h = hypandrium. ig = inner gonite. og = outer gonite. pg = proctiger. ss = surstylus. N. capilo differs from other species of subgenus Neurochaeta in its less depressed thorax, undifferentiated anterior sternopleural bristles, distinct curved anal crossvein PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 49 closing anal cell and presence of sclerotized vein 6 beyond this cell, narrow forwardly bent cercus in O" and small inconspicuous cercus in 9 . Inversa Group Inner vertical bristle present; suborbital bristle curved outwards; prelabrum moderately developed; palpus well developed but rather short; prementum rather narrow; thorax moderately depressed; humeral callus rather elongate, but without slender anterior prolongation; propleuron with well developed convex callus defined be- low by a shallow depression; scutellum with transverse series of 4 to 8 posterior bristles between major pair; upper margin of subscutellum inflexed throughout; anterior ster- nopleural bristles in a definite series, black, tapered; hypopleuron bare; prosternum more or less lanceolate, but slightly variable; metasternum with at most few minute set- ulae which are difficult to distinguish from pubescence; mid coxae separated by dis- tinctly less than width of each coxa; mid femur with short anteroventral bristle, without ventral comb; tarsal claws moderately long and strong, slightly shorter than terminal segment on each tarsus; vein 5 extending beyond discal cell; anal cell reduced, not dis- tinctly closed distally; vein 6 undeveloped beyond anal cell; cerci of both sexes exserted, broad and plate-like. Woodley (1982) introduced the inversa group to include Neurochaeta inversa McAlpine, N. sabroskyt Woodley, and N. macalpine: Woodley. With the addition of three more species, this concept is raised to a subgenus, and only the Australian N. inversa remains in the redefined znversa group. Neurochaeta inversa (Fig. 1) is the best known species in the family and has been the subject of biological and ecological studies (McAlpine, 1978: Shaw, Cantrell, and Houston, 1982; Shaw and Cantrell, 1983a, 1983b). Magnifica Group n.gr. Inner vertical bristle absent; suborbital bristle vestigial or absent; prelabrum moderately developed; palpus well developed but rather short; prementum as in inversa group, but more strongly bristled; thorax greatly depressed; humeral callus rather com- pact, not anteriorly prolonged; propleural callus large, strongly convex, as deep as humeral callus, separated from ventral section of propleuron by a deep groove; anterior notopleural bristle closer to posterior notopleural than to humeral callus, situated higher above notopleural suture than is posterior notopleural; scutellum with transverse series of 7 to 14 short posterior bristles between major pair; upper margin of subscutel- lum strongly inflexed throughout; anterior sternopleural bristles in an irregular series, black, tapered; hypopleuron with few fine setulae; prosternum very broad; metaster- num strongly setulose; mid coxae separated by much more than width of each coxa; mid femur without well defined ventral comb; tarsal claws moderately long and strong, those of mid tarsus shorter; costal armature simplified; vein 5 extending well beyond discal cell; anal cell reduced; vein 6 not visible beyond anal cell; cerci of both sexes exserted, broad and plate-like. The only known species of the group is much the largest of neurochaetids. It is the only neurochaetid yet known from New Guinea. Neurochaeta magnifica n.sp. (Figs 15, 23-25) o@ Q. With the group characters given above, otherwise agreeing with description given for N. inversa (see McAlpine, 1978) except as indicated below. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 50 STUDIES IN UPSIDE-DOWN FLIES. I Figs 23-25. Neurochaeta magnifica. 23. Prosternum. 24. Dorsal sclerites of male postabdomen (not compressed). 25. Head. bs = basisternum. ep = epandrium. fs = furcasternum. s8 = sternite 8. t6 = tergite 6. Coloration: head brown-black with grey pruinescence; face mid-brown; eye, after relaxation of freshly dried specimen, rich coppery purple; antenna brown; prelabrum and palpus dark brown; hairs and bristles on head and its appendages black, except for pale hairs on labella and antennal segment 3. Thorax coloured as in N. inversa. Mid coxa blackish; other coxae largely brown; femora brown-black, often darker distally; tibiae and tarsi black. Wing smoky brown, becoming paler posteriorly and sub-basally, but without paler zones except in immature specimens; alula dark. Haltere dark brown. Abdomen almost entirely dark brown (tergites 1-9 in 9); cercus creamy with silvery pruinescence. Head more depressed than in other species of genus; eye longer than high, with posterior margin sinuate-oblique; postfrons more narrowed posteriorly than in other species, with narrow, deep median groove in front of anterior ocellus; cheek c. 0.06-0.11 of height of eye; postvertical bristles apparently represented by a pair of convergent, approximated setulae far behind vertex; ocellar bristles very small; anterior and posterior fronto-orbital bristles markedly shorter than intermediate one, the anterior ones less medially inclined than in other species (sometimes almost parallel). Antennal sockets more widely separated than in N. inversa and N. parviceps, only slightly more so than in N. capilo. ¢ ¢ PROC. LINN. SOC. N.S.W., 110 (1), (1987) 198 D. K. MCALPINE 51 Thorax flatter than in other species (including N. parviceps), anteriorly narrowed; scutellum more broadly flattened than in other species, shorter than in parviceps group. Fore coxa not keeled; fore femur stouter than in other species, greatly swollen basally, slightly concave on anterodorsal surface; other femora more elongate than in other species, the hind one much the larger; mid femur with several short anterior to antero- ventral bristles; hind femur distally with one anterior and one or 2 posterodorsal bristles; mid and, particularly, hind tibiae, and mid tarsus unusually slender. Costal section on second costal cell with 2 rows of distally lengthening black setulae but without additional dorsal setulae just before subcostal break; costal section on marginal cell with 3 rows of closely placed black setulae for most of length, without widely spaced antero- dorsal and anteroventral bristles (weak in N. znversa); veins 3 and 4 distally convergent; costal section between veins 3 and 4 three times as long as anterior crossvein, which is well basad of middle of discal cell; costal index 5.2-5.9; vein 4 index 1.3-1.4. Abdomen. Sternites broader than in other species, in 9 sternites 4-6 as broad as tergites, in O’ 4and 5 slightly narrower. 0’ postabdomen: segment 6 with large, slightly asymmetrical tergite, with well developed symmetrically placed and formed ventral sclerite on each side, the left member of this pair apparently representing sternite 6, each of these sclerites with one long bristle and one or 2 setulae; sternite 8 reduced, asymmetrical, setulose; epandrium somewhat reduced, with 4 fine setulae only, which are not mounted on tubercles; surstylus strongly but gradually expanded basally, with basal plate less developed than in other species of subgenus, particularly on anterior side, with several setulae, mainly on posterior side well before apex, which is blunt but not thickened; outer gonite short, broad, tapered, strongly setulose; inner gonite broad, not lobed, subtruncate, incurved, with 3 setigerous tubercles on outer surface; distiphallus only moderately slender, with well developed triangular teeth, much less numerous than in N. parviceps, on about basal third; cercus broadly ovate-rhomboid, with bristles much shorter than in N. inversa and N. parviceps. Dimensions: total length, O& 5.2mm, 9 4.5-5.8mm; length of thorax, & 1.9mm, 9 1.5-2.1mm; length of wing, 0’ 4.9mm, Q 3.9-5.0mm; length of distiphallus c. 1.5mm. Distribution: Papua New Guinea — Western Highlands Province. HolotypeQ2: Mur Mur Pass, 2760m, on Pandanus leaf, 15.11.1986 (Australian Museum), J. W. Ismay. Paratypes: same data as holotype, 1 ©’, 2 9, Australian Museum, 1 9 British Museum (Natural History), 1 9 National Museum of Natural History, Washington, 3 Q Department of Primary Industry, Konedobu. Notes: N. magnifica differs from all other known neurochaetids in its large size and broad prosternum. It differs from all other species of the genus Neurochaeta in the absence of the inner vertical bristles, the posterodorsal displacement of the anterior notopleural bristle, and in having the hind femur much longer than the thorax. Parviceps Group n.gr. Inner vertical bristle present; suborbital bristle curved downwards; prelabrum much reduced; palpus very short; prementum broad; thorax much depressed; humeral callus markedly elongate, with slender anterior prolongation; propleuron without callus; scutellum without hairs or additional bristles between major pair of bristles; upper margin of subscutellum not inflexed; anterior sternopleural bristles in curved series, yellowish, scarcely tapered; prosternum narrow-linear; metasternum with well developed setulae; mid coxae separated by at least width of each coxa; mid femur usually with short anteroventral bristle before middle and loose ventral comb basally; tarsal claws short, on hind tarsus not more than % as long as terminal tarsal segment; PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 52 STUDIES IN UPSIDE-DOWN FLIES. I vein 5 not extending beyond discal cell, which has its posterodistal angle rounded off; anal cell reduced, not distinctly closed distally; vein 6 undeveloped beyond anal cell; cerci of both sexes exserted, broad and plate-like. This group includes 3 known species, viz. N. sabroskyi, N. parviceps, and N. macalpine.. All are oriental. Neurochaeta sabroskyi Woodley Neurochaeta sabrosky: Woodley, 1982: 211-212, fig. 1. This species has the characters of the parviceps group, but the thorax is more slender and the wing more elongate than in other species of the group. It is distinguishable from N. parviceps and N. macalpinei by the characters given in the key. Distribution: Philippines — Mindanao. Material examined. holotype 9 (Museum of Comparative Zoology, Harvard). Neurochaeta macalpinei Woodley Neurochaeta macalpine: Woodley, 1982: 212-214, fig. 2. This species has the characters of the parviceps group, but differs from the other two species in its paler coloration and shortening of certain bristles, including the scutellars, though the humeral and both notopleural bristles are long. Male postabdominal characters resemble those of N. parviceps. The pair of long bristles on sternite 8 is not quite as long; the surstylus is not capitate at the apex; the outer gonite is more slender and falcate; the inner gonite has a very short acute outer lobe and a more elongate curved inner lobe bearing two preapical setulae; the aedeagus has cuticular teeth almost as well developed as those of N. parviceps; there is one particu- larly strong bristle on each cercus, the other bristles being mostly much smaller. Distribution: East Malaysia — Sabah. Material examined: holotype © and 19 paratypes (Australian Museum). Neurochaeta parviceps n.sp. (Figs 5, 11, 16, 22, 26-28) o Q. Resembling WN. inversa and agreeing with description given for that species (McAlpine, 1978), except as indicated below and in the characters given for the parviceps group. Coloration: Anterior margin of postfrons, face, and anterior part of cheek creamy white; cheek bristles predominantly yellowish (© ) or blackish (9 ). Antenna, including bristles, entirely pale yellowish except for brown distal part of arista; palpus brown, paler distally. Thorax blackish with very thin greyish pruinescence; postscutellum silvery, as in N. inversa. Legs darker than in N. znversa; mid and hind femora very narrowly yellowish at bases; fore tibia yellowish, with brownish zone before middle; mid and hind tibiae black, with narrowly yellowish apices. Wing coloured as in N. inversa, but with pale sub-basal zone smaller and less distinct; haltere brown basally, with whitish capitellum. Head more anteroposteriorly compressed than in N. inversa, with postfrons longer in proportion to face and cheek narrower; postfrons concave, with linear median channel; inner vertical bristle notably shorter than outer vertical; anterior fronto-orbital bristle somewhat closer to ptilinal suture than to next fronto-orbital; suborbital bristle curved downwards. Bristles and setulae on antennal segment 2 much stronger than in N. inversa. Prelabrum vestigial; palpus very short; prementum broader than in other species. Thorax remarkably broad and flat, narrowed anteriorly; humeral callus remarkably prolonged anteriorly; scutellum a little longer than in N. inversa, with only one pair of PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 333 Figs 26-29. Neurochaeta parviceps. 26. Head. 27. Wing (ventral). 28. Dorsal sclerites of male postabdomen. 29. Neurochaeta inversa, dorsal sclerites of male postabdomen. ep = epandrium. so = suborbital bristle. s8 = sternite 8. t6 = tergite 6. marginal bristles and no hairs between the major bristle pair; pleura much excavated between anterior notopleural bristle and fore coxa; anterior sternopleural bristles 5 to 8 in a curved series, directed ventrally; sclerotized prosternal plate even narrower than in N. inversa; metasternum with numerous distinct setulae. Fore coxa more compressed than in WN. znversa, with distinct anterodorsal keel on distal part; mid coxae separated by slightly more than width of one coxa (separated by about 0.7 of that width in WN. znversa); hind femur somewhat curved on basal part (straight in N. znversa), with oblique series of short anterodorsal to dorsal bristles well before middle, a short anteroventral bristle just beyond middle, and a long dorsal bristle just beyond this. Costa, between humeral cross- vein and subcostal break, with setulae of anterodorsal and anteroventral series all black, short, moderately stout and sharp, both series only slightly increasing in size distally, in addition one thick, blunt, yellowish dorsal setula at apex of section before subcostal break and a similar dorsal setula (occasionally 2 setulae) a little basad of this; vein 4 distally attenuated, strongly converging with vein 3; costal index 4.2-4.8; vein 4 index LIS Decee Abdomen: © postabdomen: sternite 8 much reduced, glabrous, transversely elongate; epandrium somewhat reduced, with pair of long bristles and pair of fine PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 54 STUDIES IN UPSIDE-DOWN FLIES. I setulae arising from tubercles on posterior margin; surstylus rather slender beyond basal plate, almost straight, slightly thickened at apex, with minute setulae near apex only; outer gonite not as slender as in N. znversa, but more so than in WN. capilo, glabrous; inner gonite bilobed apically, but much more narrowly so than in N. capilo, with 2 setulae at apex of anterior lobe; distiphallus elongate, but stouter than in N. znversa and N. capilo and much shorter than in the latter, with rather numerous strong, short teeth on surface, instead of the weak denticles of those species; cercus somewhat as in N. inversa. Cercus of Q somewhat as in N. inversa, with one posteriorly directed bristle longer and more prominent than other bristles. Dimensions: total length, O 2.9-3.2mm, Q 3.0-3.2mm; length of thorax, o 1.3- 14mm, Q 1.3-1.4mm; length of wing, O 2.4-2.6mm, Q 2.5-2.6mm; length of distiphallus c. 0.9mm. Distribution: West Malaysia — Selangor. Holotype o&: Old Bentong Pass, E of Gombak, 16.iv.1985 (Austra!ian Museum), K. C. Khoo and D. K. McAlpine. Paratypes: same locality, 14-16.1v.1985, 22 0, 2 9, Australian Museum, 1 o’, 1 Q, British Museum (Natural History), 1 0’ Canadian National Collection, Ottawa, 1 o National Museum of Natural History, Washington, 1 ©, Természettudomanyi Muzeum, Budapest. Notes: N. parviceps differs from the other two species of the parviceps group in having the head significantly narrower than the thorax. It further differs from N. macalpinez in the darker femora and longer dorsocentral and scutellar bristles, and differs from N. sabroskyt in the entirely pale bristles on antennal segment 2 and the broadly flattened thorax. GENUS NOTHOASTEIA MALLOCH Nothoasteia Malloch, 1936: 259. Type species N. platycephala Malloch. The following are the essential characters of the genus so far as they are understood at present. The detailed description given for N. clausa no doubt includes some charac- ters which will be found to be generic rather than specific when a better range of material becomes available. Head much depressed, with reduced bristling; outer vertical bristle developed; no anterior incurved fronto-orbital bristle distinguishable; eye horizontally elongate, its surface setulose except towards posterior extremity; ocelli rather widely spaced, lateral ones at posterior limit of postfrons; antennal segment 2 less enlarged and cucullate than in Neurochaeta, more as in Anthoclusia; thoracic bristling reduced; humeral callus com- pact, rounded, very prominent, with few setulae and a weak posteriorly directed bristle; metasternum elongate, bare, separated from hypopleuron on each side by membrane of hind coxal cavity (broader than long and continuously sclerotized with hypopleuron in front of coxal cavity in Neurochaeta); mid leg remarkably small; hind coxae short, inserted on each side of metasternum (inserted behind metasternum in Neurochaeta); tarsi without functional claws; costa unbroken; second basal cell confluent with discal cell, separate from first basal cell; anal cell wide open distally; vein 6 strongly sclerotized, though not reaching margin; haltere with elongate capitellum. Only two specimens of Nothoasteia are known, but these are from remote localities in southern Queensland and southwestern Australia respectively. This, together with the fact that the two specimens represent two species, suggests the possibility that the genus has a wide distribution over Australia and includes numerous species. Perhaps, when some clues as to the ecology of the insects become available, significant data on PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 3D) distribution and species diversity may be obtained. The characters given in the above description have been confirmed for both species of the genus. Key to species of Nothoasteia 1 Discal cell widely open; hind tibia pale yellowish, faintly browned EUDL@ Ayaan Ae ane cc Meant Wh aaens teetGN ay Sh agetilli ahve uae es abe yy platycephala — Discal cell closed distally, narrower than submarginal cell; hind tibia brownish on much of length with isolated dark brown apical zone ... clausa Nothoasteia clausa n.sp. (Figs 6, 12, 30) Q . Coloration. Head with its appendages and macrotrichia pale yellowish; postfrons largely brownish, with pale pruinescence and pale yellowish anterior margin; ocellar region darker. Thorax brownish, darkest dorsally, with yellowish macrotrichia; mesoscutum and scutellum densely pruinescent; humeral callus and parts of pleura yellowish, the latter somewhat shining. Legs yellowish; hind femur with ill-defined brownish distal zone; terminal segment only of each tarsus brownish. Wing with yellow- ish veins and unpigmented membrane. Haltere pale yellow. Abdomen yellowish brown, paler ventrally. Head apparently slightly wider than long; outer vertical bristle short but distinct, somewhat proclinate; inner vertical bristle perhaps represented by a short proclinate setula slightly in front of a straight line drawn from lateral ocellus to postvertical bristle; ocellar bristles represented by pair of fine proclinate, subparallel setulae; frontal orbit with a series of several short, mostly proclinate setulae and, a little in front of middle, one moderately fine bristle sloped somewhat forwards and outwards; median part of postfrons with proclinate setulae, mostly on anterior half; frontal lunule concealed; face shallow and apparently rather narrow (collapsed); cheek region rather broad, largely facing ventrally in anterior part, much expanding posteriorly, with rather numerous long proclinate setulae anteriorly, among which some possibly represent peristomal bristles; eye with numerous procurved setulae, which are little developed on posterior part. Antenna with segment 1 short and simple; segment 2 setulose, particularly on inner side, with deep dorsolateral slit distally, with distal surface facing ventrally; seg- ment 3 probably oval and apically rounded (collapsed), without long hairs; arista minutely pubescent throughout, more densely so towards base, with basal segments (4 and 5) apparently very short (4, if present, not visible without preparation). Prelabrum rather weak; palpus moderately short; proboscis rather short with moderately developed prementum; labella separated from one another by a deep anterior slit (labella fused in Neurochaeta), each anteriorly truncate, with about 9 dentate pseudotracheae. (TET ae =) as Fig. 30. Nothoasteia clausa, wing (dorsal). PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 56 STUDIES IN UPSIDE-DOWN FLIES. I Thorax slender, rather convex above, ventrally rather planate, so that much of sternopleural region faces ventrally; 2 weak notopleural bristles present, anterior one slightly further from notopleural suture than is posterior one; a much smaller bristle or strong setula in approximately presutural or posthumeral position; mesocutum apparently without further differentiated bristles, with sparsely scattered short setulae, those on anterior part directed posteriorly (as usual), most of those on posterior half directed anteriorly; scutellar suture deeply and narrowly incised; scutellum very short, flat-topped, with free margin narrowly convex and with evenly rounded outline, with bristles reduced to 3 pairs of small, medially inclined, submarginal setulae, those of apical pair not widely separated; postnotum shallower than in most Neurochaeta spp., almost evenly convexly prominent; propleuron without evident bristles or setulae; mesopleuron with several relatively long bristles on posterior half, not concentrated near posterior margin; upper posterior sternopleural bristle represented by a minute setula; upper anterior sternopleural bristles fairly long, about 6 in a horizontal series, with apices apparently directed posteriorly; sternopleuron otherwise with scattered setulae; prosternal plate (basisternum) absent, but anteriorly angular prothoracic furcasternum distinct as in Neurochaeta (the latter perhaps mistaken for prosternal plate by Malloch, 1936). Legs having greater disparity of size than in most species of Neuwro- chaeta; fore coxa with several anterior setulae, but no anterior basal bristle; femora, tibiae, and tarsi rather irregularly haired; fore femur with a few dorsal bristles; tarsi depressed distally, with normal pulvilli; claws not visible under high magnification; one or 2 short ciliate hairs terminating fore and mid tarsi perhaps representing claw vestiges. Wing surface entirely densely microtrichose; costa unbroken, thickened apically between veins 3 and 4, on section bordering costal cell with a series of regular slightly spaced setulae, beyond vein 1 with numerous setulae or hairs, which are very diverse and irregular in size and position, on last 2 sections (beyond vein 2) with an irregular series of ventral setulae, and on dorsal surface near centre of terminal section with one rather strong setula; humeral crossvein ill defined; subcosta almost totally fused with stem vein and vein 1, visible ventrally as a ridge along anterior margin of this vein- complex, which is expanded where it joins costa; marginal cell narrow, in part no wider than thickness of vein 2; veins 3 and 4 apically convergent; anterior crossvein very short so that veins 3 and 4 are approximated at this point; discal cell rather narrow, almost parallel-sided on much of its length; terminal section of vein 5 almost as long as penulti- mate section of vein 4; posterior marginal fringe rather long, not reaching vicinity of alula; alula moderately developed, subacute; costal index c. 4.9; vein 4 index 3.4. Haltere with remarkably elongate capitellum. Abdomen slender apparently subcylindrical (not depressed as in Neurochaeta); tergites up to tergite 6 large but lightly sclerotized, with reduced bristling; pleural mem- brane thrown into telescopic folds at junctions of segments 2 to 6, as 1s intersegmental membrane; sternite 1 vestigial; sternites 2 to 6 rather broad, the more anterior ones weakly sclerotized; sternite 7 completely divided longitudinally on median line, more strongly setulose than preceding sternites; cercus not distinctly visible, probably very small. Dimensions: total length 2.2mm; length of thorax 0.74mm; length of wing 2.0mm. Distribution: Western Australia — southwest district south of Perth. Holotype 2 (unique): Yaragil 4P Catchment, via Dwellingup, 20-27.x1.1980 (Aus- tralian Museum), A. C. Postle. Malaise trap. Notes: M. R. Gray, who knows the vicinity of the type locality, informs me that it is likely to be in dry sclerophyll forest. This new species is conspicuously different from N. platycephala in venation, as the discal cell is closed and veins 4 and 5 are subparallel beyond the anterior crossvein, only PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE Dy commencing to diverge distad from the discal crossvein. ‘The shape of the marginal cell also differs, being narrowest in its mid region in N. clausa. The above description has some shortcomings due to the condition of the unique type, which has also prevented presentation of a satisfactory set of illustrations. Because the cuticle is lightly sclerotized, the head has collapsed; also the thorax is damaged by the pin. I have carefully examined the displaced sections of the thoracic cuticle on the right side and believe my account of the thoracic chaetotaxy to be accurate. Nothoasteia platycephala Malloch Nothoasteia platycephala Malloch, 1936: 259-260, figs. 1, 2, 2a. The following are supplementary notes on the unsexed holotype. Its condition does not enable production of a formal description. Coloration: Postfrons apparently with pale yellow orbital margins more differen- tiated than in N. clausa. Mesoscutum (at least anteriorly) blackish, much darker than in N. clausa. Legs pale yellowish; hind femur with very diffuse pale brownish distal zone; hind tibia pale creamy, with indistinct pale brownish suffusion on distal fifth only; each tarsus with slightly browned distal segment. Wing: Marginal cell narrow, but no narrower near middle than it is distally; vein 2 shorter than in N. clausa; veins 4 and 5 diverging throughout, quite strongly so on region just beyond anterior crossvein; discal crossvein absent, no trace or indication of its position remaining. Material examined: Holotype, unsexed, ‘Brisbane, Qld’, no other original data; ‘damaged in transit. F. H. Taylor. The head, with damaged antennae, is glued by its ventral surface to the card and there is some glue on the postfrons. Much of the thorax is destroyed but the anterodorsal part and the metasternal region are visible. Detached legs and both wings are glued to the card. Only the base of the abdomen remains. GENERA AND SPECIES OF NEUROCHAETIDAE Genus Anthoclusia Hennig, 1965 “A. gephyrea Hennig, 1965. Palaearctic (Eocene-Oligocene) A. remotinervis Hennig, 1969. Palaearctic (Eocene-Oligocene) Genus Neurochaeta McAlpine, 1978 Subgenus Neurocytta n. sg. N. prisca McAlpine, 1978. Afrotropical: Zimbabwe Subgenus Neurotexis n. sg. N. stuckenberge McAlpine, 1978. Afrotropical: Madagascar Subgenus Neurochaeta s. str. N. capilo n. sp. Oriental: W. Malaysia N. inversa McAlpine, 1978. Australasian: Australia N. magnifica n. sp. Australasian: N.E. New Guinea N. sabrosky: Woodley, 1982. Oriental: Philippines N. parviceps n. sp. Oriental: W. Malaysia N. macalpine: Woodley, 1982. Oriental: E. Malaysia Genus Nothoasteia Malloch, 1936 N. clausan. sp. Australasian: Western Australia N. platycephala Malloch, 1936. Australasian: Queensland PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 58 STUDIES IN UPSIDE-DOWN FLIES. I ACKNOWLEDGEMENTS I am particularly indebted to K. C. Khoo for assistance with fieldwork both in Australia and Malaysia. J. W. Ismay collected and forwarded the material of Neurochaeta magnifica. M. Robinson, B. G. Duckworth, D. S. Kent, and A. Bowman prepared most of the illustrations. D. J. Bickel critically read the manuscript. N. E. Woodley arranged for loan of the type of Neurochaeta sabroskyt. References CoL_eEss, D. H., and MCALPINE, D. K., 1970. — Chapter 34. Diptera. In The insects of Australia: 656-740. Melbourne: Melbourne University Press. CorRYNDON, D. H., and SavaGE, R. J., 1973. — The origin and affinities of African mammal faunas. Jn HuGuHEs, N. F., (ed.): Organisms and continents through time: 121-135. London: Palaeontological Association. CRAMPTON, G. C., 1942. — The external morphology of the Diptera. Bull. Conn. geol. nat. Hist. Surv. 64: 10-165. GRIFFITHS, G. C. D., 1972. — The phylogenetic classification of Diptera Schizophora with special reference to the structure of the male postabdomen. The Hague: W. Junk. 340 pp. Harpy, G. H., 1950. — On the articulating scutellum in Diptera. Ent. mon. Mag. 86: 230. HENNIG, W., 1958. — Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandtschafts- beziehungen. Beztr. Ent. 8: 505-688. —., 1965. — Die Acalyptratae des Baltischen Bernsteins und ihre Bedeutung fur die Erforschung der phylogenetischen Entwicklung dieser Diptera-Gruppe. Stuttgart. Beitr. Naturk. 145: 215 pp. ——., 1969. — Neue Ubersicht tiber die aus dem Baltischen Bernstein bekannten Acalyptratae. Stuttgart. Beitr. Naturk. 209: 42pp. MCALPINE, D. K. 1973. — The Australian Platystomatidae (Diptera, Schizophora) with a revision of five genera. Mem. Aust. Mus. 15: 256pp. ——., 1978. — Description and biology of a new genus of flies representing a new family (Diptera, Schizophora, Neurochaetidae). Ann. Natal Mus. 23: 273-295. MALLOCH, J. R., 1936. — Notes on Australian Diptera. XXXVI. Proc. Linn. Soc. N.S.W. 51: 259-261. SHAW, D. E., and CANTRELL, B. K., 1983a. — A study of pollination of Alocasia macrorrhiza (L.) G. Don (Araceae) in southeast Queensland. Proc. Linn. Soc. N.S.W. 106: 323-335. , and , 1983b. — Further notes on seed set in Alocasia macrorrhiza (Araceae) and occurrence of Neuro- chaeta inversa (Diptera: Neurochaetidae) in Queensland. Qd Nat. 24: 71-75. , and , and HousTon, K. J., 1982. — Neurochaeta inversa McAlpine (Diptera: Neurochaetidae) and seed set in Alocasta macrorrhiza (L.) G. Don (Araceae) in southeast Queensland. Proc. Linn. Soc. N.S.W. 106: 67-82. Woop Ley, N. E., 1982. — Two new species of Neurochaeta McAlpine (Diptera: Neurochaetidae), with notes on cladistic relationships within the genus. Mem. ent. Soc. Wash. 10: 211-218. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 Studies in Upside-down Flies (Diptera: Neurochaetidae) Part II. Biology, Adaptations, and specific mating Mechanisms DAVID K. MCALPINE McALPINE, D. K. Studies in upside-down flies (Diptera: Neurochaetidae). (Part II. Biology, adaptation, and specific mating mechanisms. Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 59-82. Field observations and apparent host plants (families Araceae, Musaceae, Pandanaceae, and Zingiberaceae) are recorded for some species. Pollen-feeding is recorded for adult Neurochaeta inversa. Aspects of adult morphology and behaviour are considered in relation to adaptation, in particular to running (including running back- wards), infrequent flight, living in crevices, adult longevity, specialization and reduc- tion of options, ecological advantages of ‘upside-down’ behaviour, and diversity of male postabdominal morphology (including evolution of the protandrium). Male post- abdominal diversity in general is here placed in the category of specific mating mechanisms. This common but little understood concept is compared critically with the premating isolating mechanism concept and the specific mate recognition system concept. The legitimacy of seeking Darwinian explanations for morphological specializations is discussed. David K. McAlpine, The Australian Museum, Box A285 Sydney South, Australia 2000; manuscript recerved 21 May 1986, accepted for publication 22 July 1987. INTRODUCTION Part I of this study (McAlpine, 1988) dealt with the systematics and phylogeny of the Neurochaetidae, including the genera Neurochaeta, Nothoastera and the extinct Antho- clusia. Some information on biology, behaviour, and ecology of the Australian Neuro- chaeta inversa McAlpine has been given by McAlpine (1978), Shaw, Cantrell, and Houston (1982), and Shaw and Cantrell (1983). Neurochaeta inversa and the East Malaysian N. macalpine: Woodley (1982) are both associated with the araceous plant Alocasia macrorrhiza (L.) G. Don. Apparent host plants belonging in other families are here recorded. These are all large monocotyledons, perhaps all providing some kind of phytotelma in which the larvae may be expected to live. It is probable that adaptation to the host microhabitat has influenced some of the morphological peculiarities and diversity of adult Neurochaetidae, but variation in other characters, particularly those of the male postabdomen, is paralleled in other dipterous families and may be a by-product of the speciation process. FIELD OBSERVATIONS (WEST MALAYSIA) Neurochaetids were observed by K. C. Khoo and the author in rain forest along the Old Bentong Pass road between Gombak village and Genting Highlands turnoff on 14- 16 April 1985. Adults of Neurochaeta parviceps were found in small numbers on peduncles of the club-shaped young inflorescences of wild ginger, Zingzber spectabile Griff. Several other species of ginger (Zingiberaceae) were flowering in the area, but no neurochaetids could be found on them. These flies moved in the characteristic upside-down mode of the genus (see McAlpine, 1978), very rapidly when pressed, and their elusive movements made them difficult to capture with an aspirator, though aspiration proved the most PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 60 STUDIES IN UPSIDE-DOWN FLIES. II effective method of capture. They sometimes sheltered in the small cavities of the stem bracts, but never in the floral bracts which were always brim-full of rain water. When disturbed they eventually ran, or more rarely flew, to other parts of the plant. If persis- tently pursued they sometimes left the plant altogether. A mating pair of N. parviceps was seen by Khoo on a Zingiber plant. In this habitat some specimens of Neurochaeta capilo were also collected. These were consistently on the leaf blades and petioles of wild banana (Musa sp., Musaceae). A few specimens of N. parviceps also found on the banana were possibly recently disturbed from an adjacent ginger plant. Only certain plants of Musa in the areas appeared attractive to N. capilo, and there was evidence of repeated return of probably the same individual to such plants. One of these attractive plants supported numerous specimens of Formicosep- sis, otherwise scarce in the area, and there was frequent non-aggressive contact between these and Neurochaeta. OBSERVATIONS ON NEUROCHAETA MAGNIFICA IN PAPUA NEW GUINEA J. W. Ismay reports as follows (zn /:tt.): ‘I was collecting at Mur Mur Pass in the Tomba Gap at approximately 143°59’E, 5°50’S [2760m].... The area had been partly cleared but some young Pandanus were left. The Neurochaeta were seen on the inside of upright leaves of Pandanus, walking up and down with the head always downwards. They ranged to the tops of the leaves. The silver markings on the back of the head and behind the scutellum were conspicuous against their dark coloration. Some were caught by sweeping a net against the leaves, but most, when approached, ran down the leaf base and were pooted. They were at least as fast as N. inversa, which I have taken in Queensland’ HOST PLANTS All specimens of Neurochaeta parviceps were found on or near Zingiber spectabile. At the time only young inflorescences were present, none having reached anthesis. These young inflorescences terminate in a cluster of bracts, each of which remains full of rain- water, because of the almost daily rainfall and low evaporation rate. Several kinds of dipterous larvae were found in this liquid from three sampled inflorescences, but none of these could be identified as Neurochaeta from comparison with the known third-instar larva of N. inversa, though some very small, probably first-instar cyclorrhaphous larvae were found. I do not regard this as strong evidence that the larvae of N. parviceps do not live in the water trapped in the bracts of Zzngiber; on the contrary I think that at a later stage of development this is most likely to be the larval habitat. The sample of N. parviceps obtained by Khoo and me has the unexpected sex ratio of 31 males to 3 females. The biological significance of this is not apparent. The strong attraction of adults of Neurochaeta capilo to plants of wild banana (Musa sp.) is established by observations recorded above. If this species is as closely associated with a host species in all stages as is N. inversa, then Musa would seem to be the larval host, but those plants on which the flies were found had closely appressed petiole bases, leaving no axillary cavities, and no inflorescences. At the time the adults were collected, only a very small percentage of Musa plants had inflorescences and these appeared to have no actual or potential water-holding parts. Musa plants are, however, often reported to provide phytotelmata. The identity of the Pandanus host of Neurochaeta magnifica is uncertain. Ismay was in- formed by a villager that the plants were ‘karuka, a name used in Papua New Guinea to designate Pandanus spp. useful for production of thatching and matting, and also for their edible fruits. In highland areas of Papua New Guinea, Pandanus jiulianetti: and P. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 61 brosimos are the principal ‘karuka’ species and their fruits are an important local food source (Stone, 1982). P. brostmos occurs in the Tomba vicinity close to the type locality of Neurochaeta magnifica (Stone, 1982: figs 6, 7) and may be the host of this fly, but this is only one of about 66 species of Pandanus known from New Guinea. Stone also states: “he pandan leaf axil is of considerable interest. It is usually stocked by the infall of detritus from above, usually retains water to the extent of a cupful or more, or a thick solution of decaying debris, and may incorporate dying fragments of the endo-axillary rootlet system’ Among the invertebrates living in this habitat he mentions larvae of cyclorrhaphous flies. Thus this appears to be a likely habitat for the larvae of Neurochaeta magnifica. However, Ismay (zn /itt.) points out: ‘Since the [karuka] palms are 10-30m high and very spiny, few entomologists collect from them. Also, they cannot be tampered with — damage to karuka 1s acommon cause of tribal conflict in the Highlands’. FEEDING IN NEUROCHAETA INVERSA At Mount Tenison Woods, D’Aguilar Range, near Brisbane, Queensland, on 411.1983, K. C. Khoo and I observed inflorescences of Alocasia macrorrhiza being visited by three species of insects, all taking pollen from the spadix. These were Neurochaeta in- versa, Trigona sp. (apparently 7’ carbonara Smith, det. E. Exley, a native social bee) and Apis mellifera LL. The last species was in the smallest numbers, but tended to disturb or disperse the others when present. Clearly, higher concentrations of A. mellifera would have inhibited seriously the activities of the other two species. We observed numbers of WN. znversa adults repeatedly licking the surface of the male section of the spadix and apparently ingesting pollen. On a number of occasions the flies were seen to approach the hind leg of a Trgona and actively lick at the pollen load. The Trigona, in each case, attempted to withdraw its leg or move away, but the fly often followed to some extent. While Alocasia pollen provides a high protein food source for N. inversa, this food is available only during the summer flowering period of the plant. My previous con- jecture, that the flies feed on various substances collected on the large leaves of Alocasia, is supported by further observations (Border Ranges National Park, near Kyogle, N.S.W., 3.1v.1987). Periods of running activity on leaf surfaces were interrupted at in- tervals when the flies began licking at spots of unidentified substances on the leaf sur- face. When drops of diluted orange-marmalade were smeared in their paths, the flies stopped to feed on it. Thus the leaf-surface activity is to be interpreted as foraging. LOCOMOTION AND LEG STRUCTURE I have described for N. inversa (McAlpine, 1978) the habit of running rapidly back- wards and forwards with constant head-downwards orientation while on a vertical sur- face. This habit is now also recorded for a further four species of the subgenus Neurochaeta, viz. N. parviceps and N. capilo (observations by K. C. Khoo and author), N. macalpine (observations reported to author by J. Frazier and D. Clyne, noted Woodley, 1982), and N. magnifica (Ismay’s observations). Almost certainly the fifth species of the subgenus, N. sabroskyi, has the same habit. The speed and the erratic nature of running are remarkable in subgenus Neurochaeta for such small insects. I have observed a specimen of N. inversa to run at an average speed of about 3.7cm/sec for a period of 30sec, during which time the direction of movement was reversed about 50 times. Under similar conditions an active specimen of Stenomicra sp. of similar size moved at about 1.2cm/sec with only about 6 fairly abrupt reversals of direction in 30sec. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 62 STUDIES IN UPSIDE-DOWN FLIES. II The specialized running mechanism of Neurochaeta appears to have certain advan- tages in: (1) avoiding predators, (2) finding scarce food substances by covering a large area of plant surface in a short time, (3) seeking out small apertures (e.g. for escape from plant cavities), (4) the searching out of conspecific individuals in aggregation and perhaps in sexual activity. Differentiation of the leg proportions is characteristic of the family. These are particularly expressed in the relative size of the femora and the general description is as follows: fore femur short and stout; mid femur short and slender; hind femur long and moderately stout; tibia of each leg shorter than femur. The precise proportions of the femora vary between the species as shown in Table 1. The fossil Anthoclusia gephyrea has the least differentiation in femoral length, according to the scale drawings of Hennig (1965: figs 251, 252), and this appears to be the most plesiomorphic condition known in the family. Neurochaeta magnifica and the species of Nothoasteza, on the other hand, have the greatest differentiation. The reduction of the mid legs in Nothoasteia is reminiscent of that of wingless males of the hymenopterous family Agaonidae. It seems logical to relate the unusual leg proportions of neurochaetid flies to the mode of locomotion which is characteristic of all observed species, even though present knowledge does not explain the mechanistic aspects of this relationship. TABLE 1 Relative lengths of femora in neurochaetids Ratios fore femur: mid femur: hind femur A. gephyrea OF, 18 Tale 14s Q.- Ne. capilo Oy, ME ALS Nets} ©) teil 28 IL-9) Ne. inversa @7, Je hile LY) Q. 1: 1.1: 1.9 Ne. magnifica Or, ile N23 Bod Q. 1: 1.2: 2.6 Ne. sabroskyt oO. - ®. 1: 1.0: 1.9 Ne. parviceps Oy te walle ALG) Oe 118 als 3) Ne. macalpiner ors 18 i08 1k} Q. 1: 1.0: 1.8 No. clausa oO. - 9. 1:0.7: 2.1 No. platycephala ?. 1: 0.8: 1.9 Flies of the genus Nothoastera are remarkable and perhaps unique among the Schizophora in the absence of anything in the nature of a tarsal claw. Because tarsal claws are so generally present in Diptera, it is certain that they fulfil a function in their biology which cannot normally be dispensed with. Presumably the primary function is clinging to surfaces, which are either rough or sufficiently soft for an impression to be made by the sharp claw apex. One might, then, infer that Nothoastera normally lives on surfaces which offer no such opportunity for gripping with claws, such as a hard, smooth surface or a loose powdery one. The paired pulvilli are well developed on all tarsi of Nothoasteia, as in other neurochaetids. These are pads of sticky hairs which enable most flies to cling to and walk on smooth surfaces, even if the surface is vertical or facing downwards (e.g. a window pane or ceiling). They do not function on wet surfaces (so far as known). If non-sticky, these pads could aid walking on a powdery surface. While absence of functional claws may seem to set Nothoasteia apart from the rest of the Neurochaetidae, there is variation in claw size between species of the subgenus Neurochaeta. Also there may be variation in claw size in the one individual, those of the smaller mid legs often being slightly smaller than those of the large hind legs. Neurochaeta capilo and N. parviceps are flies of similar size, but the hind tarsal claws of the former are at least twice as long as those of the latter species. The other species of the parviceps group PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 63 have claws of similar proportions to those of N. parviceps, but N. inversa has claws inter- mediate in size between those of the capilo and parviceps groups. The strong, moderate-sized claws of Neurochaeta inversa are similar to those of periscelidids and many other small acalyptrate flies, and probably approximate to the plesiomorphic condition for the Neurochaetidae. ‘The elongate, needle-like condition in N. capilo and the shortened condition of the claws in the parviceps group appear to be apomorphies which have developed in opposite directions. Claw reduction in Nothoasteva is clearly independent of and convergent with that in the parviceps group. The adapta- tional significance of this reduction may become clearer when studies are made of living Nothoasteva. RUNNING BACKWARDS The orientation of the hairs and bristles on the dorsal surface of flies with apices directed posteriorly is apparently an adaptation to walking (and probably to flying) for- wards with minimal resistance. Though several (perhaps all) species of Newrochaeta walk backwards as much as they walk forwards (and probably for much longer periods than they fly forwards), the orientation of most hairs and bristles remains as in strictly forward-walking flies, though a few of the dorsal bristles are sometimes nearly erect. These longer bristles would tend to shield the posterior mesoscutal hairs from contacting any obstacle dorsal to the insect. By contrast the hairs on the median region of the posterior half of the mesoscutum in Nothoasteia clausa are directed forwards, those on the anterior part being largely directed backwards; most bristles are quite short and there are no long posteriorly directed ones. In the absence of behavioural records of this rare fly, this condition suggests that the backward locomotion may be at least as important as forward locomotion in Nothoasteia. Vhe legs of Nothoasteia are similar to those of some apomorphic types found in Neurochaeta, e.g. in N. parviceps, particularly in the long hind femora and reduction in size of the mid legs; also the body-form is strongly depressed, and reduction of the prosternal plate has extended to complete loss. These features suggest behavioural similarity in the two groups. Because Nothoasteia has not acquired apomorphic wing-venational characters present in the groundplan of subgenus Neuro- chaeta, it is evident that the extreme developments of the legs and body-form have been acquired independently in the two groups. As the developments are mostly present in a less elaborated state in such plesiomorphic neurochaetids as Anthoclusia and subgenus Neurocytta, it 1s not surprising that similar states of elaboration should have been achieved in different lineages. Concentration of organs of vision on the anterior end in insects is an obvious adaptation to forward locomotion, perhaps particularly to forward flight as non-flying cursorial insects (e.g. worker ants) generally have reduced eyes. Because, in subgenus Neurochaeta, running backwards is a more frequent occupation than forward flight, one might expect some modification of vision in connection with this behaviour. In all the species of the genus Neurochaeta the eyes are obliquely elongate. Thus, though total area of the eye is not great, there is a larger number of ommatidia facing dorsally than in other cursorial flies (such as phorids, and certain sphaerocerids). In living examples of N. inversa and N. parviceps, there is a characteristic concave-backed profile resulting from the head being held away from the substrate through dorsal flexion at the neck. This unusual position contrasts with that of dried specimens, and careful examination shows that several ommatidia would then provide some vision in a posterior to posterodorsal direction. The convex posterior extremity of the eye in N. magnifica could also increase posterior vision. PROC. LINN. SOG. N.S.W,, 110 (1), (1987) 1988 64 STUDIES IN UPSIDE-DOWN FLIES. II Previously (McAlpine, 1983) I recorded observations indicating that the aulacigas- trid flies Nemo centriseta McAlpine and Nemo kentae McAlpine walk consistently forward, while WN. corticeus McAlpine and N. phaeotylos McAlpine often walk backwards and for- wards with abrupt reversals of direction at short intervals. I have recently examined the eyes of these four species, and find that the posterodorsal margin of the eye in the two lat- ter species is extended slightly further on to the posterior surface of the head and has the marginal ommatidia directed slightly more in a posterior direction than in the former pair of species. Firm conclusions as to this apparent connection between behaviour and structure require more observations on Nemo spp. than those yet made. Except for N. centriseta, the observations have been few, and the morphological difference between the two species pairs is small. However, if such a connection is proved for Nemo, this would strengthen the expectation of comparable adaptations in Neurochaeta, where backward motion is a more significant element of behaviour. In neurochaetids the parts projecting furthest posteriorly, and therefore those which usually make first contact with an obstacle in running backwards, are the wing tips and the tip of the abdomen. In the subgenus Neurochaeta and in Nothoasteia the wing tips have special hairs or setulae which could be tactile. In the former they form a small, compact group at the apex of vein 3; in Nothoasteza these setulae are not in such a com- pact group, and the short section of the costa bearing them, between veins 3 and 4, is remarkably thickened. This thickening could be a strengthening device in a part subject to battering when the insect runs backwards. In most species of the subgenus Neurochaeta the cerci of both sexes are broadened, exserted, and fringed with long setulae. The possibility of these structures acting as tactile organs and buffers for running backwards could explain why the cerci, which usually have evolved along different lines in each sex, have here evolved the same apo- morphic condition in both sexes. N. capilo is the only species of the subgenus Neurochaeta with cerci short (in female) or not posteriorly prominent (in male), although it is capable of running backwards at speed. This species 1s apparently a sister group to the rest of the subgenus, and has probably never acquired these apparently adaptive attributes, despite an equally long history of running backwards. This condition of the cerci is perhaps explained by the insect having larger wings, a further plesiomorphic character. Nothoasteia also lacks modifications of the cerci or other apical abdominal parts. In Neurochaeta capilo and Nothoasteva the greater size of the wings relative to the abdomen renders it less probable that the abdomen would make first contact with an obstacle. Perhaps also N. capilo lives in a more open habitat than N. inversa, N. parviceps, and N. macalpinei, and is thus less likely to run into objects. Examples of adaptation for running backwards as well as forwards in other animals seem to occur mainly in types living in tunnels or burrows, where it can be advan- tageous to reverse direction of locomotion without turning the body. Moles of the genus Talpa (Mammalia: Insectivora) are reported to run as fast backwards as forwards (Boon- song and McNeely, 1977). The fur of moles is soft and velvety, and can lie in any direc- tion, enabling the animals to go backwards and forwards in a burrow without the grain of the fur giving resistance. These facts prompt comparison with Nothoastera and raise the questions: Does Nothoasteza live in tunnels or burrows? Can its dorsal thoracic setulae have their direction reversed, like moles’ fur, or is their position, as described above, per- manent? In Yalpa micrura Hodgson the tail is much reduced, but acts as a sensory organ when it is running backwards. This compares with the posterior sensory organs of Nothoastera and subgenus Neurochaeta. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 65 FAST FORWARD LOCOMOTION In view of the presence of apparent adaptations for rapid backward movement, comparable adaptations for forward movement in neurochaetids should also be con- sidered, as forward running appears to be just as frequent and rapid as backward run- ning. As mentioned above, forward vision is well developed. Forwardly directed vibrissae are present in Neurochaeta, but the anterior parts most liable to the effects of col- lision are the antennae. Antennal segment 3 is presumably furnished with the usual complex and delicate array of sense organs found in higher Diptera, including chemoreceptors. The rather long hairs on the anterior surface of this segment would afford some protection to these organs in Neurochaeta, and segment 3 is particularly well protected from physical contact with surrounding objects by the large, cucullate seg- ment 2, which, in subgenus Neurochaeta, bears strong anterior bristles. The arista projects further forwards than other parts of the antenna and could serve to sense an imminent collision rather than to shield physically other parts from contact. In species of subgenus Neurochaeta, which, so far as known, tend to flex the head upwards, the anteriorly inclined median ocellus would be unable to receive stimulation from a light source directly in front of the insect, were it not for the fact that the post- frons has a median channel leading to the space between the antennae. In some species, e.g. N. magnifica and N. parviceps it is scarcely wider than the ocellus, very well defined, and commences immediately in front of the ocellus. INFREQUENT FLIGHT Observations on species of the subgenus Neurochaeta seem to indicate that running Is a more frequently used escape mechanism than flying and that flight is an infrequently used form of locomotion. Nevertheless all species for which we have field observations can fly (i.e. all species except N. sabroskyz), and flight is probably necessary for dispersal to new host plants. N. macalpine: shows reduction of the wing area, which is probably an indication of the lesser importance of flight in its biology. Comparison of the wings in N. macalpinet and N. parviceps suggest that the latter also has undergone slight reduction of the wing and that the loss of the free distal part of vein 5 is an element of this reduction (also occurring in a third species, N. sabroskyz, with slightly longer but narrower wing). N. parviceps often seemed most reluctant to fly in the field, probably more so than N. capilo which lacks these indications of wing-reduction (though field observations on the latter species were restricted to few individuals). N. macalpine: not only shows the greatest wing reduction of any known Neurochaeta species, but also the most marked shortening of many of the bristles, notably the fronto- orbitals, dorsocentrals, and scutellars, though the notopleurals are quite long. There seems a possibility that the bristle-shortening is an adaptation to infrequent flight, though the importance of bristles in flight is not well understood. On the other hand Nothostera species show a much greater bristle reduction without apparent reduction of wing area. This is not just a reduction in length, but an overall reduction in size and number of the bristles, and,in parts, also a reduction of the hairs. The case of Neurochaeta macalpinei is reminiscent of the genera Baeopterus (Coelopidae) and Calycopteryx (Micro- pezidae), both flightless examples with shortened bristles in families of normally actively flying forms. LIVING IN CREVICES I have already concluded (McAlpine, 1978) that in Neurochaeta inversa the dorso- ventrally compressed body is an adaptation to moving through narrow spaces, such as are provided by the host plant, Alocasia. This ability has been confirmed on two PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 66 STUDIES IN UPSIDE-DOWN FLIES. II occasions when several adults of N. znversa escaped from a collecting jar by way of the thread of the screw-top, which had an imperfect inner seal. By comparison, N. capilo is much less depressed. Perhaps, if /Zusa is the only host of this species, then there is less necessity and opportunity for creeping into narrow spaces. One individual, apparently of N. capilo, when pursued, was seen to shelter in the channel on the adaxial side of the banana petiole, but this was a relatively open and capacious hollow. Neurochaeta parviceps has a more strongly depressed body than other Neurochaeta species which I have observed in the field, though in N. macalpinei it is almost as depressed and in N. magnifica more so. N. parviceps has been seen to shelter in the narrow and shallow spaces in the axils of the bracts on the peduncles of young inflorescences of Zingiber spec- tabile. A reduced depth of the thoracic region by comparison with N. inversa is partly achieved through reduction in size of the mid coxae and their migration from near the median line to a more lateral position on the thorax. This results in a greater ventral exposure of the metasternum (which in N. znversa tends to be concealed by the mid trochanters). In the parviceps and magnifica groups the ventrally directed anterior sterno- pleural bristles and the setulae on the metasternum are particularly well developed, in response, I believe, to the need for feeling the substrate in an insect in which the thorax is held unusually close to it. The thoracic pleura of N. parviceps and to some extent those of related species (including N. inversa) have, laterally to the fore coxa, a marked ventrally facing hollow, which appears to enable relatively free movement of the laterally splayed fore legs, without their occupying space between the thorax and the substrate. The size and apparent resting position of the fore coxa, as evidenced from dried material, varies among species of Neurochaetidae, and these attributes relate to the variations in structure of the prothoracic furcasternum. In Neurochaeta capilo the fur- casternum is not prominent and is covered by the more or less distally contiguous, bulky fore coxae. The coxae appear to work largely below the sternal region of the thorax, occupying a significant part of the depth of the insect. In N. parviceps the broad, flat fur- casternum rather widely separates the more laterally placed fore coxae. The coxae are less bulky than in N. capilo, compressed, keeled, and apparently adapted for movement within the pleural hollow without occupying much space below the thorax. In N. inversa the condition is intermediate between that of N. capilo and N. parviceps. The furcaster- num is rather narrowly convex and the coxae, though decidedly separated at rest, are generally less so than in N. parviceps. The condition in N. macalpiner and N. sabroskyi 1s similar to that of N. parviceps, but in the former the rather small coxae are neither compressed nor keeled. In Neurochaeta magnifica the structure and co-adaptation of these prothoracic parts are quite different from the above types. The problem of the working of the forelegs under a remarkably shallow thorax has been solved by a migration of the coxa almost to the lateral extremity of the thorax, the section of the pleura above it being almost vertical and exceedingly shallow to accommodate the coxa. With the fore coxae extremely dis- tant from each other and the prothoracic pleura quite limited to the lateral surfaces of the thorax, there remains a ventral prothoracic surface much broader and more open than in other species. The furcasternum is thus even broader than in N. parviceps, and the very broad prosternal plate is unlike that of any other neurochaetid. In Nothoastera clausa the condition of the fore coxae most resembles that of Neurochaeta macalpinei, but there is no pleural hollow. In contrast to the mid coxa, mobility of the fore coxa is essential to the operation of the leg in locomotion; hence the quite different nature of its specialization. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 67 Figs 1, 2. Metasternum and surrounding parts of Neurochaeta spp. 1. N. inversa. 2. N. parviceps. c2 = mid coxa. c3 = hind coxa. s3 = metasternum. tr2 = mid trochanter The erect dorsal bristles on the wing and scutellum in the subgenus Neurochaeta are probably important tactile organs when the insect moves under a low ceiling. There is a similarity here to suberect apical scutellar bristles of the genus Nemo (family Aulacigas- tridae, see McAlpine, 1983). At least some species of Nemo shelter under bark, a habitat which provides a low ceiling. The plant association is unknown for Neurochaeta sabroskyi, but its proportions suggest a slightly different direction of specialization from that of related species. It seems probable that it is a sister species to NV. parviceps, and out-group comparison both for the species pair and the collective group tends to confirm its derivation from broadly depressed forms. Yet N. sabroskyi has a much more slender body and wings. Although the fore coxae somewhat resemble those of N. parviceps, the pleural hollow is less marked, because narrowing of the mesoscutum allows less overhang of the pleural region. Hence we appear to have an adaptation pattern which has moved from wide, shallow crevices to something like pin-holes, or, more probably, when the accommodation of the legs is considered, in grooves which are both narrow and shallow. APPEARANCE AND POSSIBLE MIMICRY On exhibiting living material of Newrochaeta to laymen, a typical comment is some- thing like: ‘Are they flies? They look more like ants. ‘This reaction is presumably the result of several visible features. The size and slenderness of these insects are likely to be attributed to certain familiar domestic ants (e.g. species of genera Jredomyrmex and Tech- nomyrmex) rather than to such familiar flies as domestic calliphorids and muscids. These attributes are, however, typical of numerous flies of the superfamily Asteioidea (in which Neurochaetidae are currently placed), and alone cannot be considered as evidence for mimicry of small ants by these flies. I shall consider some other characters which appear to support the view that some neurochaetids are Batesian mimics of small dolichoderine ants. Adults of Neurochaeta parviceps and N. inversa have a general blackish body coloration relieved by a paler zone at the anterior end of the abdomen. The idea that this coloration may be of some adaptive significance receives support from the fact that it is emphasized by the pale sub-basal zone of the wings, when they are flexed over the abdomen, and also by the coloration of the halteres. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 68 STUDIES IN UPSIDE-DOWN FLIES. II In N. parviceps and N. inversa in the subgenus Neurochaeta the halteres are bicoloured, with pedicels brownish and capitella creamy white. With the haltere directed posteriorly in the resting position, the dark pedicel lies against the dark posterior part of the thorax, and the whitish capitellum lies against the pale base of the abdomen. The wings being translucent, the coloration of the haltere reinforces the insect’s longitudinal sequence of colour zones. The coincidence of the distal parts of both wings over the abdomen gives the wings a low degree of visibility. The total impression, then, is of a slender, dark, wingless insect with a posterior part or gaster somewhat separated from the rest of the body, ie. the appearance of a dark-coloured dolichoderine worker ant about 3 mm in length. Comparison of Neurochaeta species with temperate Australian species of the micropezid genus Metopochetus tends to convince me that wing pattern in both is an element of Batesian mimicry of ants. Metopochetus species of the taxonomically uneluci- dated M. terminalis (Walker) complex are almost certainly mimics of ants of the genus Leptomyrmex (see Colless and McAlpine, 1970: fig. 34.29A, where incorrectly given as M. tenuipes). The larger Metopochetus compressus (Walker) is probably a mimic of aggressive stinging ants of the genus Myrmecia. These micropezid species have a complex wing pattern reinforced by superposition of the wings over the abdomen when at rest. The wings thus visually tend to lose their identity and give the impression of a basally narrowed, segmented abdomen. A closely related undescribed species of Metopochetus from Lord Howe Island has no wing pigmentation. Apparently there are no suitable ants to serve as models on this oceanic island. Several other micropezids, e.g. Mimegralla contingens (Walker) in northern Australia, New Guinea etc. and Jaeniaptera spp. in Brazil, have a wing pattern similar to that of Metopochetus spp., though not closely related. Again I believe this convergence in pattern is best explained as due to ant-mimicry. Neurochaeta species (e.g. N. inversa, see McAlpine, 1978: fig. 3) have a simpler wing pattern than the ant-mimicking Metopochetus species, probably because the smaller size of the former renders detailed representation of abdominal segmentation unnecessary. Whether or not one is convinced of my theory of ant mimicry by Neurochaeta species, the evidence that colour pattern in these insects is adaptive is strongly supported by the fact that colour pattern of different parts of the insect is co-ordinated not only in such species as Neurochaeta parviceps and N. inversa as explained above, but also in other species with different schemes of coloration. N. sabrosky: has the capitellum of the haltere tawny, unlike other species of the genus, and also an almost uniformly brown-tinged wing membrane, and the anterior abdominal tergites brownish. N. macalpinei, on the other hand, has larger pale areas on the body than other species, paler legs, paler bristles, entirely creamy white halteres, and a larger pale sub-basal wing zone. The co- ordination of colour in wing, haltere and abdomen is apparent, and there is a possibility (but at present no evidence) that this species is a mimic of a small pale-coloured ant. In collecting the type series of N. macalpinei, Clyne and Frazier obtained, in association with the flies, a bug of the heteropterous family Anthocoridae. This is of similar size and coloration to the flies and could be a mimic of the same ant mimicked by them or even a mimic of the flies themselves. There is also a possibility that it is a predator of Neurochaeta. The zigzag movements of Neurochaeta spp. are not particularly ant-like, though ants often do run quickly. This movement could, however, serve to display and reinforce the ant-like signal already learned by the predator. A further feature observed in Neurochaeta parviceps and N. inversa, which is unusual for the higher Diptera and which increases resemblance to an ant, is the prognathous position of the head due to dorsal flexion at the neck in the living insect. The apparently short, high head of dried material is not ant-like, by comparison. The upward tilting of PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 69 the head has been interpreted above as possibly aiding vision in a posterior direction, but I see no reason why it may not also aid in protective mimicry. Ants, including small dolichoderines, have relatively long conspicuous antennae. Neurochaetids have not been observed to compensate visually for this lack by an appropriate position or motion of the fore legs, as has been observed in a number of dipterous mimics of Hymenoptera (McAlpine, 1973: 9-10). On the other hand, general impression rather than precision of detail is probably more important in mimics of such small size. Muimicry of ants by other insects and spiders is a common and well established phenomenon (Wickler, 1968; author’s numerous observations, and numerous other references in literature). It has occasionally been doubted that ants are suitable models for Batesian mimicry because they are preyed upon heavily by insectivorous vertebrates, the predators through which visual selection is most likely to operate. On the other hand I believe it probable that ants which contain acrid chemicals are likely to bring about predator satiation at relatively low levels of predation, at least in non-specialist preda- tors. The fact that ant numbers are often very large increases both the probability of pre- dator satiation and the probability that the predator will learn the visual signal. Ants with such a powerful sting as Myrmecia naturally produce a cautious approach in an experienced predator, which is likely to increase the chance of escape for a mimic. It is interesting to note that the Baltic amber fossil neurochaetid Anthoclusia gephyrea Hennig (1965: fig. 244) has a wing pattern somewhat resembling that of Neurochaeta inversa. Uhis suggests the possibility that neurochaetids have been mimics of ants for as long as 40 million years. ADULT LONGEVITY From what is known of the life-cycle of Neurochaeta inversa in New South Wales, over- wintering female adults would need to live for about 6 months in order to find an ovi- position site and may in fact live even longer. Over-wintering males seem to live about as long as females, from my observations, and mating appears not to take place till an oviposition site is available (see McAlpine, 1978). Though I am not aware of any longevity experiments for such small flies in the field, it is probable that N. znversa adults have rather exceptional longevity for their size class. Perhaps such behavioural peculiarities as the extreme agility in running, the tendency to seek shelter, the devices which protect the wings from battering, the infrequent use of flight, and ingestion of such protein-rich food as pollen are adaptations to prolonged survival. It would be interesting to compare longevity, behaviour, and relation to the seasonal cycles of the host plants of the equatorial Malaysian species with those of N. znversa. SPECIALIZATION AND REDUCTION OF OPTIONS As usual in cases of extreme specialization, the locomotory behaviour in Neurochaeta seems to have narrowed the range of biological options. Thus fast running is carried out almost exclusively with a strong forward or backward component, and, for reasons still unexplained, the insect is unable to remain in any position except that with the body axis vertical while on a vertical surface. It is possible that even this restriction on orienta- tion is adaptive, as discussed below. The elongate body in N. inversa appears to be remarkably rigid. When the insect feeds from a leaf surface, the head is not flexed from the neck, but the body remains straight as its anterior end is sloped towards the substrate. Observation on a captive specimen of N. parviceps indicated a restriction in loco- motion due to rigidity of the body. The insect was placed in a stoppered cylindrical glass PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 70 STUDIES IN UPSIDE-DOWN FLIES. II specimen tube c. 25mm by 50mm. The tube was inverted to give the container a glass ceiling. As expected, the insect ran up and down on the vertical glass wall with the body vertical and the head downwards. It repeatedly ran against the glass ceiling but each time it ran down again after contact. It was apparently unable to cross over the junction of the two surfaces at right angles to one another, and was reluctant to turn the body to make this transition possible. Eventually, after much contact with the ceiling, the body was turned horizontally and the transition to the ceiling readily made by walking side- ways. Clearly inability to bend the body adequately, perhaps combined with shortness of the mid legs, prevented passing from the vertical to the horizontal surface when starting from the normal vertical body-position. No such difficulty existed for the insect in pass- ing back from the ceiling to the wall, because, on the horizontal ceiling, there was no tendency towards a constant orientation of the body which could have prevented it approaching side-on to the vertical wall. It is doubtful if the artificial situation just described simulates any frequently encountered natural situation. Flies on more mature inflorescences than those observed by us would frequently encounter the almost horizontal upper margins of bracts, and passing over this margin directly from the outer to the inner surface could pose a problem similar to that just described. One would expect, however, that any frequent, naturally occurring situation would not provide such difficulties in problem-solving. Field observations of N. parviceps in these situations would be interesting. WHY UPSIDE-DOWN? Given the advantages which may be conferred by the habit of running rapidly backwards and forwards in zigzags, the question remains as to why this habit has evolved in association with a head-down instead of an unrestricted or head-up orienta- tion of the body. It has been noted above that Neurochaeta species (and also Nothoasteia species) show apparent adaptations for running backwards, that these modifications are not uniform throughout the Neurochaetidae, and that the neurochaetid body-plan, like that of other Diptera, is primarily adapted for forward movement. In particular the eyes and other important sense-organs, which provide information as to conditions likely to be encoun- tered during locomotion, are located on the head. It seems advantageous, then, for the neurochaetid fly to orient itself with the head pointing in the direction in which obstacles or dangers are most likely to be found. The cavities in which these flies shelter, e.g. the axils of bracts and hollowed petioles, and the spathal cavity of Alocasza, all open upwards, their closed lower ends forming an obstacle which, on repeated contact would eventually damage the wings. In the special case of phytotelmata, the opening is always upwards, and it would seem safest to approach the liquid surface, a potential sticky trap, head first. I have often noticed how moisture drops in glass vials tend to trap small flies by the wings. Thus, running wings-first towards the liquid surface could be particularly dangerous. Some likely predators of Neurochaeta such as scincid lizards and frogs shelter in plant axils. Therefore running downwards into an axil head-first would seem more advan- tageous for flies than the reverse. Facing downwards while resting within the cavity does not, however, seem an advantageous position for watching for predators, and the neuro- chaetid habit of emerging from shelter backwards seems incautious. There is, however, a degree of rear vision and emerging backwards may not be particularly risky. The running is likely to be rapid and, as usual, evasive during emergence. A predator would need to be watching a particular axil carefully to be able to take advantage of a backward emergence. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 71 ee . S6 xX Ay ae, aa Voir if 79) Ye ae J / Figs 3-6. Protandrograms of schizophorous Diptera, showing segments in front of male genital segment as if split along median ventral line and spread flat. 3. Coelopa frigida (Fabricius). 4. Neurochaeta prisca (recon- structed from notes and sketches). 5. N. capilo. 6. N. magnifica. s5-s8 = sternites 5-8. t5-t8 = tergites 5-8. x = supernumerary sclerite. It is interesting to note that flies of the genus Stenomicra (family Periscelididae or Stenomicridae), which often share the habitat of Neurochaeta species and are probably also associated with phytotelmata, maintain a consistent head-upwards orientation. But Stenomicra are not known to shelter in axils, they run less rapidly than the observed Neuro- chaeta species, and they do not move backwards and forwards in a zigzag path (McAlpine, 1978). I postulate that the head-downwards orientation in the Neurochaetidae evolved during the early stages of development of zigzag running. In view of apparent adapta- tions for running backwards in Nothoasteza, this kind of orientation may be expected to occur in all known living neurochaetids, and possibly occurred in the forms known from fossils. But it is conceivable that this orientation did not become rigidly stereotyped, as in the subgenus Neurochaeta, until the flies acquired habitat preferences which brought them into frequent contact with phytotelmata. DIVERSITY IN MALE POSTABDOMINAL STRUCTURES The few known species of the genus Neurochaeta show a great diversity of structure in the segments behind segment 5 of the male abdomen. The male is unknown in subgenus Neurotexts. Variation in sternites 6-8 consists of degree of reduction of these sclerites. Sternites 6 and 7 are well developed in Neurochaeta (Neurocytta) prisca (Fig. 4), where they are strongly asymmetrically placed. In this form the dorsal sternite 8 is also well developed. Subgenus Neurocytta thus has the most plesiomorphic condition of these sternites known in the genus, as may be seen from the resemblance to the primitive schizophoran Coelopa (Fig. 3). PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 72 STUDIES IN UPSIDE-DOWN FLIES. II The remaining known species of the genus (subgenus Neurochaeta) show a trend towards reduction of sternites 6-8 and towards symmetry of these segments. N. capilo (Fig. 5) has sternites 6 and 7 much reduced but still discernible on the left side, while sternite 8 remains large and setulose. N. macalpine: and N. parviceps have these segments externally approximately symmetrical with a small sclerite below each lateral margin of tergite 6, and sternite 8 reduced to a narrow glabrous transverse strip. N. inversa resem- bles the above species in the symmetry of segment 6 but sternite 8 is absent. I previously (McAlpine, 1978) interpreted a dorsal sclerite of N. inversa as probably sternite 8, but comparison with other species of the subgenus not then available, indicates that the sclerite in question is the epandrium. The protandrial structure of Neurochaeta magnifica is of special interest, most of its characters being strongly apomorphic. In contrast with that of other species of subgenus Neurochaeta, tergite 6, though unreduced, is asymmetrical, as is the reduced sternite 8. Ventrally segment 6 bears a symmetrical pair of lateral plates, each separated from the tergite by the pleural membrane as is the sternite of the previous segment. Each plate bears macrotrichia which are better developed than those of sternite 5. Were the phylogenetic derivation of N. magnifica not reasonably clear, one would probably in- terpret this structure as the result of median desclerotization of a large, primarily sym- metrical sternite 6 resembling the preabdominal sternites (Fig. 6). However, the apparent facts that the inversa group 1s the sister group of the magnifica group, that these together form the sister group of the parviceps group, and that the three above groups together form the sister group of the capzlo group, render such an interpretation 1m- plausible. The series of protandrograms (Figs 3-6) illustrates the direction of evolution of these structures in the genus Neurochaeta. Only the left ventral sclerite of segment 6 in N. magnifica can be the homologue of sternite 6, its mirror image on the right side being a secondary sclerite. This condition is somewhat paralleled in Fannia (Griffiths, 1972) and Borboroides (McAlpine, 1985), where structures present only on the left side in the ances- tral form have been mirrored by new structures on the right side, with resultant sym- metry. In NV. magnifica, however, 1t appears that evolution of segment 6 1s in the process of incorporation into the preabdomen, as a continuation of the sclerotization process would restore a full, symmetrical sclerite simulating a typical preabdominal sternite. This kind of process may well be the clue to the possession of an apparent symmetrical, ventral sternite 6 in males of such flies as Waterhousera (family Heleomyzidae, see McAlpine, 1985). It has long appeared to me from study of other families that the groundplan condition of the Schizophora includes strongly asymmetrical protandrial sclerites as in Coelopidae (Crampton, 1942: fig. 12H), and that symmetrical conditions of the protandrium are all secondarily derived. As protandrial morphology has figured prominently in arguments on phylogeny and superfamily classification (see particularly Griffiths, 1972), the correct evolutionary interpretation 1s important. Some aspects of the functional changes in the evolution of the neurochaetid pro- tandrium are easily explained. The asymmetry of the sclerites was apparently attained in a remote Mesozoic ancestor through spiral displacement in the process of circum- version of the genital segment (Crampton, 1942). This spiral arrangement of the tergites and sternites in segments 5 to 8 is preserved in modern Coelopidae with almost diagrammatic clarity (Fig. 3). With slight modification, this pattern occurs in the plesio- morphic neurochaetid Neurochaeta (Neurocytta) prisca. N. prisca is a relatively large neuro- chaetid, and size in the groundplan of subgenus Neurochaeta had probably been reduced to no more than about 3 mm total length as exemplified by 5 of the 6 known species. Size reduction in flies, as in other animals, is often accompanied by structural simplification, possibly because of ontogenetic difficulties in producing a diversity of structures from smal] materials, or perhaps mainly because a smaller organism can function on a PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 78) simplified plan more readily than can a larger organism. In particular, a smaller body requires less skeletal support. In any case the subgenus Neurochaeta has developed a tendency towards symmetry by reduction, N. capilo (Fig. 5) and N. parviceps showing different stages of this process. Apart from the size reduction-simplification factor, the inherent instability in copulatory structures resulting from speciation processes, con- sidered below, combined with the mechanical requirements of approximate symmetry in flight, are factors which should contribute to development of symmetry. The increase in body size and consequent need for skeletal support, and for a base for additional sen- sory macrotrichia in N. magnifica has resulted in increased ventral sclerotization of seg- ment 6. But in this case there has been no tendency for this reversal of selection pressure to restore the lost asymmetrical sclerites, and resclerotization of the region has produced a new almost symmetrical pattern. In N. prisca the epandrium is well developed and the surstyli are loosely articulated with its margins. In the species of subgenus Neurochaeta the surstyli are detached from the reduced epandrium and arise each from its separate basal plate. The gonites (paired appendages of the hypandrium) consist of a single pair in N. prisca and two pairs in subgenus Neurochaeta. Like the surstyli they show specific differ- ences in shape. The aedeagus shows considerable specific variation in length, thickness, and armature in the genus Neurochaeta. Specific differences in male copulatory organs occur in most families of Diptera, sometimes being more remarkable than in the examples cited above, e.g. those in the aedeagus of the heleomyzid genus Diplogeomyza (McAlpine, 1967). The Diptera are not peculiar in this respect as such specific characters are so general in insects that taxo- nomic work on most orders normally takes these characters into account. Probably for insect groups in general it may be stated that the male copulatory organs are more con- sistently than any other organs the ones which show morphological divergence between closely related species. There is a parallel here with acoustical behaviour, though the latter is probably of less wide occurrence among insects. Highly specific genitalia characters occur in other groups of animals besides insects. They have been described in many Acari (e.g. Davis, 1968), in Diplopoda (e.g. Johns, 1964), in monogenetic trematodes (Sproston, 1946), in gastropods (e.g. Solem, 1981), in snakes (Dowling and Savage, 1960), in carnivorous marsupials (Woolley, 1982), and in rodents (e.g. Lidicker, 1968). These are probably all cases where there is or has been a possibility of the sexual stages of related species mixing with one another. There has been difference of opinion as to the reason for specific diversity in the genitalia of insects. Some have held that these specific differences constitute a ‘lock and key’ isolating mechanism (e.g. Watson, 1966). Mayr (1963: 104), while admitting that mechanical isolation (e.g. by means of genitalia difference) plays ‘a very minor role’, explains this diversity as neither adaptive nor contributing to isolation, but as a side effect of pleiotropic genes. SPECIFIC MATING MECHANISMS Previously (McAlpine, 1969) I have classified specific difference in genitalia as an element of a ‘specific mating mechanism rather than an isolating mechanism. A specific mating mechanism is defined as any genetically based device which tends to prevent mating or attempted cross-fertilization between species. Common specific mating mechanisms among insects include specific differences in time or place of courtship, differences of courtship pattern, visible appearance, chemical secretion, song, or tactile PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 74 STUDIES IN UPSIDE-DOWN FLIES. II recognition marks, and the necessarily correlated specific responses to the stimuli imposed by these. Specific mating mechanisms have often been classed as ‘premating isolating mechanisms or interpreteted as ‘reinforcement of isolation through selection. They are, however, distinct from true isolating mechanisms, in that they are the result of, not necessarily the cause of, some degree of isolation between species, and the ‘species iso- lation concept is at best a misnomer. Their magnitude, complexity, and intraspecific constancy is only to be explained through their role as specific recognition marks, con- ditioned through inviability or inferior fitness (including heterozygote disadvantage) of the hybrids which they tend to eliminate. Inferior hybrid fitness is the true cause of iso- lation (McAlpine, 1969). It is pointed out that even mechanisms of low efficiency can be produced by natural selection, and that in nature hybrids can occur between species having well developed specific mating mechanisms. However, the combined effects of specific mating mechanisms can amount to almost 100% efficiency. Many pairs of species which hybridize readily in the laboratory rarely, if ever do so in nature, even when they are sympatric (Mayr, 1963; Bock, 1984). In a study of acoustical behaviour in the so-called races of the Drosophila paulistorum complex, Bennet-Clark and Ewing (1970) found evidence suggesting that the original barrier to interbreeding 1s not a difference in song, which is only a secondary mechan- ism. The fact that sterility barriers are evolving with or even preceding differentiation of song illustrates my point that such specific mating mechanisms are not the real cause of isolation. The clear distinction between isolation and the specific mating mechanism can be seen in the following example from literature. Fisher (1958) cites recurrent hybrids between the butterflies Lzmenitis arthemis and L. astyanax occurring at low frequency in the narrow zone of overlap of these two species. A strong mating preference (specific mating mechanisms on my understanding) is said to be responsible for this low fre- quency. Apparently the hybrid lineages are eliminated after a few generations because of inferior fitness. If Fisher’s data are accurate, the gene pools of the two species are as effectively isolated as if every hybrid zygote perished. He is therefore mistaken in sug- gesting the populations to be subspecies rather than species. I would agree with Fisher that the ‘sexual preference’ is favoured by a selective process, but not that this process ‘would establish an effective isolation’, as effectively complete isolation continues even when the ‘sexual preference’ breaks down, though with some waste of activity and gametes. In a reasonably stable population the genes of the overall genotype tend to make up a highly integrated set, and it has long been known that introduction of chromosomal material of one race into that of another can cause significant changes in fitness to the carriers of the mixed genotype (e.g. Dobzhansky and Spassky, 1944). In the case of sym- patric interfertile incipient species (which have recently acquired sympatry or are the products of disruptive selection) the hybrid genotypes very generally have inferior fit- ness to the parental genotypes, or the parental populations will tend to lose their identi- ties. Hence, selection for a specific mating mechanism can occur through interaction with a related species even if the hybrids are viable and fertile. Thomas (1950) has described the intimate association of the male genital append- ages with the vaginal opening of the female during copulation in Sarcophaga (Diptera, Sarcophagidae). In this genus the form of the external genitalia shows a high degree of variability which is quite specific, and the female is known to exercise a very precise choice on males attempting to copulate. The conclusion seems inescapable that the pattern of stimulation caused by the structure and perhaps the movements of the male genitalia assist the female to make this choice. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE HS) The various male postabdominal organs which show great variation in the genus Neurochaeta seem likely to affect the process of copulation in some way, so that a specific response is possible in the female. ‘The large asymmetrically placed sternites 6 and 7 in subgenus Neurocytta, absent in subgenus Neurochaeta, seem likely to affect the articulation of the genital segment with the preabdomen and thus the mode of wielding the copu- latory organs. The paired periphallic appendages, surstyli and gonites, most probably convey tactile stimuli to the female. There can be no doubt that the great difference in length of the aedeagus, together with difference in its cuticular armature between N. capilo and N. parviceps, enables specific discrimination by females, should any premating behavioural mechanism break down. Though adults of these two species are usually separated by host-plant preference, they can occasionally occur on the same plant. Despite the views of Eberhard (1985) I find in male genitalia characters some evidence of the geographical character dislacement pattern dependent on patterns of sympatry. In the genus Pseudopomyza s.1. (Diptera, Pseudopomyzidae) the species occur- ring in Europe and that in Australia, each widely geographically isolated from others of the genus (probably for a very long period) and not very closely related to each other, have the surstyli essentially similar in shape. In New Zealand, where there are several species which must have evolved with some degree of contact with one another (Harrison, 1959) the surstyli show strong specific differences in shape from those of the above species and from each other. A similar pattern occurs in the genus Australimyza (Diptera, Carnidae), where the several New Zealand species have highly specific shapes for the surstyli, but the two species, which are geographically remote from others of the genus, in Australia and Macquarie Island respectively, have similar and relatively simple surstyli. Again, these last two do not appear closely related on the basis of other characters. In flies of the genus Euprosopia and in some other genera of Platystomatidae there is a sclerotized, capsule-like segment of the aedeagus or penis called the glans which is inserted deeply into the female genital tract in copulation. The glans often differs in size between closely related species but is of remarkably constant size within a species, despite great individual variation in body size. In Queensland, the two partially sym- patric species E. separata and E. comes are so similar that females are difficult to dis- tinguish, but males are distinguishable by the size of the glans and some other secondary sexual characters (some information in McAlpine, 1973a). In the more southern part of its range, where it is the only species of its group, E. comes apparently has a glans inter- mediate in size between that of E. separata and the more northern populations of E. comes. Despite such examples as this I find a detailed uniformity in genitalia characters over a considerable geographic range to be usual in the dipterous groups I have studied. These data seem to conflict with the pattern given by Eberhard (1985: chapter 3). In some, but not all, of Eberhard’s cases the genitalic difference between allopatrics con- cerns species within genera of considerable specific diversity and the species compared are not necessarily very closely related. Again he may have too readily dismissed the likely significance of unknown historical patterns which could often have been quite complex. The existing allopatry often may be irrelevant to the speciation process. On the other hand some of Eberhard’s arguments seem to presuppose that only coexistence of very closely related species can possibly be invoked as producing sexual selection for divergence per se, whereas copulations and sometimes hybridism in nature between species of a higher order of differentiation have been observed (e.g. in butterflies and birds). I summarize my conclusions on specific mating mechanisms (SMMs) in the following 5 statements: — PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 76 STUDIES IN UPSIDE-DOWN FLIES. II 1. SMMs arise through natural selection acting on sympatric populations of related species. 2. Production of SMMs is not part of the speciation process, though it commonly follows speciation. 3. SMMs reduce wastage of gametes and of time and energy in fruitless reproductive behaviour. 4. SMMs do not (in the long term) prevent introgression (do not cause isolation). 5. SMMsare probably generally maintained by selection. It is now necessary to consider hypotheses which are not compatible with the SMM viewpoint, but which have arisen partly through some shortcomings of the isolation- reinforcement concept. H. E. H. Paterson has often stated the view that, while recognition marks are stabilized within a species by natural selection, natural selection has not caused diver- gence in these characters between related sympatric species (the concept of the specific- mate recognition system or SMRS, see Lambert and Paterson, 1984). According to that viewpoint, divergence in characters involving recognition occurs as a result of random genetic changes in populations of small size before speciation. While the small popu- lation model of speciation has become widely accepted, taken alone it is quite incapable of explaining the widespread (in some groups quite general) phenomenon of much greater specific divergence in premating signals than in other characters. Mate recog- nition would still be an important stabilizing factor at the small population stage, though under reduced mate-choice some slightly deviant phenotypes may be accepted. As with the older speciation model based on geographic variation, an initiation of signal divergence could result, but in my view, there is no reason why, under allopatry, divergence in signal should vastly outstrip divergence in other characters. This final result is clearly directed, not random, and almost certainly depends on sympatry (some examples in genitalia characters given by McAlpine, 1969). Paterson (1978) supports his view, that the reinforcement model (thus also the specific-mating model) cannot be evolved through interaction between incipient species, by a simple mathematical demonstration. This idea, that, if there is random mating and heterozygote disadvantage in a mixture of two genotypes, the rarer geno- type will be so often absorbed into unfit heterozygotes that its extinction is assured, 1s valid under certain ideal circumstances (experiments of Harper and Lambert, 1983), but let us compare these requirements with the situation in nature. First, why is it necessary to assume random mating? Paterson accepts that evolution after allpatric separation of a small population makes some divergence in ‘SMRS' possible, and thus mating choice in subsequent sympatry need not be entirely random. It has been argued, however, that, where behavioural differences occur, they cannot alone prevent the extinction of one genotype in a closed system. Second, the development of ‘reinforcement’ or specific mating mechanisms occurs between popu- lations that have diverged to the point of speciation, not between simple mutants of the one population. Third, and perhaps most important, the model depends on a nice mixing under closed conditions of the two genotypes, as in the few laboratory experi- ments which Paterson considers ideal. In nature the patchiness of the environment renders it more likely that there would be frequent invasion and possibly return between strongholds of the two population types. After each invasion the original occupier would have lost some material from the gene pool, preferentially that most likely to have facili- tated cross mating. Any retreating invader would be similarly changed. The very experi- ments which have been rejected as irrelevant, those in which a mixed culture was artificially maintained, are those most likely to simulate a natural event. It has been pointed out that these laboratory experiments do not result in permanent fixation of PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE WA mate recognition systems in the populations, but this is scarcely surprising. The estab- lished systems or mechanisms in natural species have often been evolved over many thousands of years, and even then may be imperfect enough to allow occasional hybrids, e.g. in birds of paradise (Gilliard, 1969) and birdwing butterflies (McAlpine, 1970), both groups with obvious specific recognition marks. There is reason to believe that, in sym- patric sister species, specific mating mechanisms are maintained by selection, just as they were built up by selection. When changes in the environment bring about break- down in the true isolating mechanism of hybrid inferiority, the species tend to fuse, because the specific mating mechanism can no longer be maintained (several examples given by Mayr, 1963: chapter 6). It is not quite reasonable of Lambert and Paterson (1984), whenever there is geographic variation in signal characters, to claim that this variation cannot be in those characteristics of the signal that involve recognition. If the observed variation does not involve recognition marks, why else should it follow the geographic character- displacement pattern observed, for instance, by Littlejohn (1965)? Littlejohn’s expla- nation fits the observed facts, but the SMRS school provides no adequate explanation of observed phenomena. On the other hand, the positive aspect of the SMRS argument does explain the remarkable consistency of specific mating mechanisms within a species, which may extend throughout its geographic range if gene flow permits. Certainly, some of my above points have been considered by the anti-reinforcement school. For example, Harper and Lambert (1983) consider the effect of continued immigration of one species, but argue that continued gene flow (i.e. introgression) would probably prevent divergence of the populations. As I have previously pointed out, the divergent genomes (under the allopatric speciation model) need to have reached, prior to sympatry, a degree of divergence which ensures that each integrated genome has such superior fitness that hybrid lines are virtually certain to die out in the long term. Populations that have not so diverged are irrelevant to the speciation process, even though they may, if allopatric, be given separate specific names. Eberhard (1985) discusses Fisher’s concept of sexual selection by runaway female choice and favours this explanation for specific divergence in genitalia characters over selection for divergence under at least partial sympatry. Kirkpatrick (1982) has produced a mathematical model for this system. Eberhard argues that the other func- tion of copulation (i.e. other than essential sperm transfer) ‘is that of inducing females to receive and use sperm or, in a broad sense, courtship. The tendency for females to be selective in their sexual partners is a well known corollary of the facts of production of different numbers of gametes and the making of different kinds of investment in the off- spring by the two sexes. This principle is also significant in the specific mating mechan- ism concept. The runaway process can indeed be invoked to explain rapid change in these circumstances, but that the change is under a special kind of control is evidenced by the end result, viz. remarkable uniformity within a species and sharp interspecific difference. These almost universal phenonema do not necessarily arise as the con- sequence of ‘runaway selection’ as described particularly by Eberhard. The theoretical effects of this selection, especially under the explanations given by Eberhard (1985:72), should be a continuing instability and elaboration of detail through female choice favouring novel stimuli. Very generally, such instability does not occur in male genitalia characters. Amazingly, when Eberhard comes to consider intraspecific uniformity of genitalia as a stumbling block to his hypothesis (Eberhard, 1985: 151-153), he finds this evidence ‘difficult to evaluate’, and then all but denies its existence. There is also a weakness in the converse side of the runaway female choice argument. Why, if the process takes place quite independently in each species, does it virtually always (in the relevant animal groups) result in strong divergence in genitalic PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 78 STUDIES IN UPSIDE-DOWN FLIES. II characters? It might be expected that in the more complex systems change per se would usually result in divergence, but the divergence phenomenon is quite general for various levels of structural complexity. Some further inconsistencies occur in Eberhard’s arguments. He argues (a) that copulation is part of the courtship ritual (b) that copulation is not necessarily the final stage at which female choice can be exserted. (I can agree on both counts.) He then argues that the fact, that the poor correlation of simple, uniform (i.e. non-specific) male genitalia characters with (specific) elaborate premating behaviour, seriously weakens the species isolation hypothesis (presumably meaning for genitalia differences as distinct from courtship differences). As indicated elsewhere, I do not consider any one mechan- ism to be 100% efficient to the exclusion of other mechanisms; further there is no neces- sity 1n any case to believe that the premating mechanism was evolved before the genitalic one, and special problems in gamete wastage will favour reinforcement of recognition mechanisms at all stages in the sexual sequence. Finally, there are many cases where closely related species do occur sympatrically, are interfertile, and occasionally produce hybrids. The fact that these hybrids (and not the parent species) are eliminated means that ‘reinforcement’ selection is certainly acting. I find it just about as surprising, then, that some biologists should attempt to disprove the existence of this selection process, as that they should attempt to disprove the existence of upside-down flies. TAXONOMIC CHARACTERS AND ADAPTATION The view that genetic changes which result in taxonomic characters are generally differentially adaptive is probably widely accepted. A taxonomic character, if genuine, is generally the phenotypic expression of a number of genes, though it may not represent the whole adaptive expression of any one of its causative genes. The theory of the adap- tive nature of taxonomic difference is closely related to the specific niche theory. For both theories, support can be found in many specific examples, but a general proof is not available. Species divergence need not, however, depend on competition. It is quite possible for taxonomic difference to be adaptive without its bearing on any ecological difference between taxa. Often a taxonomic difference is adaptive to the internal environment of an organism which has itself become modified for various historic and probably adaptive reasons. The difference in chaetotaxy (bristle pattern) between such flies as Homoneura (family Lauxaniidae) and Leucophenga (family Drosophilidae) may be of this kind. The existing pattern in each group is stable, apparently because it fits the very complex physiology of the organism, although muta- tions liable to cause different patterns are often produced. Drosophilids manage best without a mesopleural bristle; lauxaniids need a mesopleural bristle (except in the special case of the Celyphinae), though flies of the two groups seem to be performing the same things. I have consistently sought Darwinian explanations for morphological change and divergence, because no established modern theory has been able to displace ‘progressive adaptation as the driving force in morphological evolution. Neither the theory of molecular drive (Dover, 1982) nor the neutral theory of molecular evolution (outlined by Kumura, 1985) is claimed to have much direct bearing on morphological change, though both are concerned with change at the molecular level. Sometimes, however, hypotheses regarding morphological developments of unknown function are put forward, and there can be a tendency to interpret these under the above categories. Vines (1982), perhaps finding difficulty in explaining specific differences as the effects of selection, suggests that molecular drive may have an effect ‘on any aspect of the PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 79 phenotype, from sexual behaviour to morphology” If this generalization is to be applied to the phenotypes here classed as specific mating mechanisms, then its protagonists must explain the curious coincidence that these phenotypes are always those which have the potential to enable specific discrimination in mating pairs. Arrow (1951), discussing evolution of enlarged mandibles and horns in Coleoptera, infers a form of evolutionary momentum when he states ‘because a process [of evolution | is long continued it is not easily discontinued’. Such ideas have been largely countered by responsible arguments (e.g. Otte and Stayman, 1979; Charlesworth, 1984). Recent interesting studies of evidence for function of diverse developments have included: stalked eyes in Diptera (McAlpine, 1979; Burkhardt and Motte, 1983), horns in Coleop- tera (Otte and Stayman, 1979; Eberhard, 1979 — Arrow’s problem!); horns in Diptera (Moulds, 1977); diverse secondary sexual developments in Diptera (elements of specific mating mechanisms, McAlpine, 1973b); shell geometry in gastropods (Signor, 1985); coloration in insects (a few examples and references in Matthews, 1976); stripes in zebras (Cloudsley-Thompson, 1984 — involving Diptera!); the tusk in male narwhals (Cetacea) (Gerson and Hickie, 1985). The last example provides support for the analogy I have drawn (McAlpine, 1976) between sinistrally spiral narwhal tusks and sinistrally spiral vibrissae in certain clusiid flies. The examples here quoted can be categorized with most of the morphological types sometimes considered inexplicable in terms of Darwinian selection. The need for alternative theories seems to be disappearing as we take a closer look at organisms in nature. In the above discussion in general I have been using the term adaptation in virtually a traditional sense, but also in the special sense of Gould and Vrba (1982). That my interpretation of usage of the morphological developments should sometimes extend these characters into the category of exaptation (using the terminology of Gould and Vrba) is for the most part improbable and not intentional. This is because I am con- sidering new developments, which are apomorphies in relation to the groundplan of the Asteioidea and often in relation to that of the Neurochaetidae or even of subgenus Neuro- chaeta. Hence the characters are not likely to be preadapted to uses other than those per- taining to the derived biological pattern of neurochaetids. The possible exception involves the prognathous position of the head in subgenus Neurochaeta, which appears to have dual usage in ant mimicry and rear vision. However, there is no reason to assume that prognathy preceded the early stages of either ant mimicry or running backwards and it may be a simultaneous adaptation to two functions. Attempts by taxonomists to interpret in functional or adaptive terms the characters used in classification are occasionally made, mainly in the more obvious categories, such as specific diversity of bills of birds (examples in Tyne and Berger, 1959). In insects, with the enormous number of taxa and consequently of taxonomic characters, few taxonomists have given time to consider the functional aspects of the characters, and lack of relevant biological data has often placed the problem beyond profitable consideration. Hlavac (1972) has given a morphological account of the prothorax of Coleoptera, relating the major structural types to locomotory mechanism and habitat adaptation. The differences in structure are to some extent the character differences for major taxa of Coleoptera. He finds that, at somewhat lower taxonomic levels convergence is common ‘and putatively unique paradaptive features’ are infrequent. This statement recalls the apparent convergence in characters relating to thoracic depression between Nothoasteia and the more advanced species of the Neurochaeta lineage. Generation of morpho-adaptive hypotheses for the Neurochaetidae is simpler than for most families of Diptera, because of the small number of taxa, the fairly clear picture PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 80 STUDIES IN UPSIDE-DOWN FLIES. II of part of the phylogeny (partly from reference to a key fossil), and some understanding of their biology. In discussing the morphology of the Neurochaetidae I have produced a series of hypotheses as to adaptive values, which are in accord with the very limited available data on the biology of the insects, or which, considered in relation to one another, make up a plausible picture of aspects of the evolutionary adaptation of neurochaetid flies. In taking my deductions somewhat into the field of speculation I have proceeded further than is usual in the field of insect physiology, but not further than is commonly accepted in the fields of palaeontology and phylogenetic systematics, where elaborate hypotheses are often produced on evidence which 1s slight or liable to more than one interpretation. As in those fields, I believe that reasonable hypotheses on morphological adaptation in insects are a step towards a more complete understanding of a little investigated but highly complex subject, and provide some indications for the direction of future work. ACKNOWLEDGEMENTS I am indebted to W. D. Hamilton, R. W. Taylor and D. J. Bickel for discussion of some of the ideas here put forward. K. C. Khoo assisted in field observations, B. Duckworth assisted with the illustrations, and J. Howarth processed the words. References ARROW, G. J., 1951. — Horned beetles, a study of the fantastic in nature. The Hague: W. Junk. BENNETCLARK, H. 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N.S.W., 110 (1), (1987) 1988 D. K. MCALPINE 81 Jouns, P. M., 1964. — The Sphaerotrichopidae (Diplopoda) of New Zealand. 1. Introduction, revision of some known species and descriptions of new species. Rec. Canterbury (N.Z.) Mus. 8: 1-49. Kimura, M., 1985. — The neutral theory of molecular evolution. New Sczent. 1464: 41-46. KIRKPATRICK, M., 1982. — Sexual selection and the evolution of female choice. Evolution 36: 1-12. LAMBERT, D. M., and PATERSON, H. E. H., 1984. — On ‘bridging the gap between race and species’: the isolation concept and an alternative. Proc. Linn. Soc. N.S.W. 107: 501-514. LipIckER, W. Z., 1968. — A phylogeny of New Guinea rodent genera based on phallic morphology. /. Mammal. 49: 609-643. LITTLEJOHN, M. J., 1965. — Premating isolation in the Hyla ewing: complex (Anura: Hylidae). Evolution 19: 234-243, MCALPINE, D. K., 1967. — The Australian species of Diplogeomyza and allied genera (Diptera, Heleo- myzidae). Proc. Linn. Soc. N.S.W. 92: 74-106. ——, 1969. — Systematic studies in Australian Platystomatidae (Diptera). London: University of London, Ph.D. thesis, unpubl. — , 1970. — Anote on the status of Ornzthoptera allotte: (Rothschild) (Lepidoptera: Papilionidae). /. Aust. ent. Soc. 9: 233-234. ——, 1973a. — The Australian Platystomatidae (Diptera, Schizophora) with a revision of five genera. Mem. Aust. Mus. 15: 256 pp. ——, 1973b. — Observations on sexual behaviour in some Australian Platystomatidae (Diptera, Schizophora). Rec. Aust. Mus. 29: 1-10. —., 1976. — Spiral vibrissae in some clusiid flies (Diptera, Schizophora). Aust. ent. Mag. 3: 75-78. —, 1978. — Description and biology of a new genus of flies representing a new family (Diptera, Schizophora, Neurochaetidae). Ann. Natal Mus. 23: 273-295. —., 1979. — Agonistic behaviour in Achzas australis (Diptera, Platystomatidae) and the significance of eye- stalks. In M.S. and N. A. BLuM, (eds) Sexual selection and reproductive competition in insects: 221-230. 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New York: Academic Press. PATERSON, H. E. H., 1978. — More evidence against speciation by reinforcement. S. Afr. J. Sct. 74: 369-371. SHAW, D. E., and CANTRELL, B. K., 1983. — Further notes on seed set in Alocasia macrorrhiza (Araceae) and occurrence of Neurochaeta inversa (Diptera: Neurochaetidae) in Queensland. Qd Nat. 24: 71-75. , and HousTon, K. J., 1982. — Neurochaeta inversa McAlpine (Diptera: Neurochaetidae) and seed set in Alocasia macrorrhiza (L.) G. Don (Araceae) in southeast Queensland. Proc. Linn. Soc. N.S.W. 106: 67-82. SIGNoR, P. W., 1985. — The role of shell geometry as a deterrent to predation in terebrid gastropods. Scz. in N. Guinea 11: 81-95. p) SOLEM, A., 1981. — Camaenid land snails from Western and Central Australia (Mollusca: Pulmonata: Camaenidae). II. Taxa from the Kimberley, Amplirhagada Iredale, 1933. Rec. West. Aust. Mus. Suppl. 11: 147-320. SPROSTON, N. G., 1946. — A synopsis of the monogenetic trematodes. Trans zool. Soc. London 25: 185-600. STONE, B. C., 1982. — New Guinea Pandanaceae: first approach to ecology and biogeography. Jn J. L. GRESSITT, (ed.), Brogeography and ecology of New Guinea 1: 401-436. THOMAS, H. T., 1950. — Field notes on the mating habits of Sarcophaga Meigen (Diptera). Proc. Roy. ent. Soc. (A) 25: 93-98. TYNE, J. VAN, and BERGER, A. J., 1959. — Fundamentals of ornithology. New York: John Wiley and Sons. VINES, G., 1982. — Molecular drive: a third force in evolution. New Scient. 96: 664-665. Watson, J. A. L., 1966. — Genital structure as an isolating mechanism in Odonata. Proc. Roy. ent. soc. Lond. (A) 41: 171-174. WICKLER, W., 1968. — Mimicry in plants and animals. New York: McGraw-Hill. WOODLEY, N. E. 1982. — Two new species of Neurochaeta McAlpine (Diptera: Neurochaetidae), with notes on cladistic relationships within the genus. Mem. ent. Soc. Wash. 10: 211-218. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 82 STUDIES IN UPSIDE-DOWN FLIES. I WooLLEY, P. A. 1982. — Phallic morphology of the Australian species of Antechinus (Dasyuridae, Marsupia- lia): a new taxonomic tool? Jn M. ARCHER, (ed.), Carnivorous marsupials 2: 767-781. Sydney: Royal Zoological Society of N.S.W. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 A new Genus and four new Species in the Family Echinasteridae (Echinodermata: Asteroidea) FRANCIS W. E. ROWE and E. LYNNE ALBERTSON ROWE, FRANCIS W. E., & ALBERTSON, E. LYNNE. A new genus and four new species in the family Echinasteridae (Echinodermata: Asteroidea). Proc. Linn. Soc. N.S.W. 110 (1), (1987) 1988: 83-100. A new genus and four new species of echinasterid asteroids are described. The species are recorded from South African, southern and southeastern Australian, Japanese and northwestern North American waters. The genus is characterized by the occurrence of a large, hyaline-tipped recurved spine at the apex of the jaw. The genus otherwise shows a close relationship with Henricia Gray (1840). EW. E. Rowe and E. L. Albertson, Division of Invertebrate Zoology, Australian Museum, Sydney, Australia, 2000; manuscript received 6 February 1987, accepted for publication 22 July 1987. INTRODUCTION During an investigation of the echinoderm fauna of New South Wales, taxonomic problems became apparent in members of the asteroid spinulosid family Echinasteri- dae. The identity of species of the genus Henvicia in southern and southeastern Australia and the description of a new species of Echinaster from New South Wales, have been reported by us elsewhere (Rowe and Albertson, 1987a, b). The present taxonomic problem deals with a geographically widespread group of new species which share in common a large recurved hyaline-tipped spine at the apex of the jaw angle. All other features are typically echinasterid. The presence of this apical tooth was identified first amongst southern and southeastern Australian specimens identified in the genus Henricia held in the Australian Museum (AM) and Museum of Victoria (MV) collections. During a visit to the United States National Museum (USNM) the senior author found two additional specimens, one from Japanese waters, the other from off Washington State, NW coast of North America. Finally two speci- mens were included amongst a collection identified as Henricia abyssalis from South African waters, on loan from the British Museum (Natural History) (BM(NH) )in London, U.K. SYSTEMATIC DESCRIPTION Family ECHINASTERIDAE Verrill (1867) Genus Odontohenricia nov. Description: An echinasterid sea star with 5 rays (rarely 6); reticulate abactinal skele- ton, plates raised centrally forming ridges which bear spinelets; marginal series distin- guishable, inferomarginal plates pronounced; papulae present abactinally, marginally and actinally; 2-3 furrow spines in vertical series; large recurved hyaline spine present at apex of each pair of oral plates, the size 3 to 4 times greater than adjacent spines. Type species: Odontohenricia endeavouri n.sp., herein designated. Etymology: odontos = (Greek) tooth, referring to the apical oral spine characterizing this genus. henricia = named after the genus Henricia, in recognition of the otherwise close morphological association of these genera. Other species included: O. clarkae n.sp., O. fisherin.sp., O. hayashii n.sp. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 84 NEW ECHINASTERID ASTEROIDS Remarks: Odontohenricia differs from all other members of the family by possessing a recurved apical spine. Otherwise the new genus appears to share most features in common with Henricia Gray, 1840, including the form of skeleton and arrangement of spines. Considering the general echinasterid features of these species, we do not believe the possession of this hyaline-tipped apical spine warrants the establishment of a new family. We do, however, consider this discrete group of species should be recognized within its own supra-specific taxon and describe a new genus accordingly. As far as we are aware, the recurved apical spine is a feature shared only with members of the valvatid family Odontasteridae. Within the Echinasteridae, Plectaster Sladen, 1889 is the only genus in which the furrow spines occur 1n a comb-shaped row along the adradial edge of the adambulacral plate (Fig. 1). We believe this genus does not belong in this family, though we decline herein to reassign it, since it is outside the scope of this paper. Fig. 1. Plectaster decanus (AM J13074): Actinal surface showing the comb-shaped rows of spines along the adradial edge of the adambulacral plates. Odontohenricia endeavouri n.sp. Figs 2A-B, 3A-B, 4A-E Henricia hyadesi, H. L. Clark, 1916: 60 (part); 1946: 148 (part) [non HZ. hyadesi (Perrier, 1891) = H. obesa (Sladen, 1889) according to Fisher, 1940: 164]. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 F. W. E. ROWE AND E. L. ALBERTSON 85 Diagnosis: Rays 5, rarely 6, R=31-83mm, r=5-14mm, R/r=5-12, inflated proximally and tapering; abactinal skeleton closely reticulate, 1-2 rarely 3 papulae per papular area; plates overlapping, irregular, ovoid to stelloid form; plates medially ridged bearing fine spinelets which taper to an acute tip, spinelets in tufts of 3-9, or in 1-3 comb-like irregular rows; marginal plates distinct, ridged medially and bear spinelets, supero- marginals and inferomarginals quadrilobed, inferomarginal plates markedly higher than wide, intermarginal plates more or less quadrilobed with small plates between, 1-3 series, actinal plates quadrilobed 1-3 series, marginal spinelets as for abactinal plates; papulae extend to actinal series with 1 papula per area; 2 (3) furrow spines in vertical series, adambulacral plates with a large cylindrical or slightly compressed sub- ambulacral spine on the adradial edge, behind which stand a pair of shorter but similar spines. Smaller serrately tipped subambulacral spines, occur across the remaining sur- face of the adambulacral plate, (these often appear to be aligned in a double row); large recurved spine at apex of oral angle, 4-5 spines occur along furrow margin of oral plates, 3-4 spines occur on actinal surface of oral plates. Materials examined: Holotype, AM J8792, off Eden, New South Wales; 1 paratype, AM J7156, Lakes Entrance, Vic.; 1 paratype, AM J5859, Bass Strait west of Babel Island, 128m; 4 paratypes, AM J20080, off Eden, N.S.W.; 1 paratype, AM J3072, Great Australian Bight, 146.3-219.5m; 4 paratypes, MV Acc. No 75-9, 38°44’S, 141°33’E, 18 miles south of Cape Nelson, Vic., 153.6m; 3 paratypes, MV Acc. No 75-9, 38°44’S, 141°33’E, 12.5 miles south of Cape Nelson, Vic., 146.3-155.4m; 1 specimen, AM J12868, 34°21°S, 151°24’E to 34°14’S, 151°28’E, northeast of Wollongong, N.S.W., 402m; 1 specimen, AM E3648, Great Australian Bight, 146-274m; 1 specimen, AM Josoype bass Strait, 1 specimen, AM) }17266, 34°43 7S loloi27E to 34°38" lolcilo/ i, N.S.W., 457m; 1 specimen, AM E3772, Great Australian Bight, 274m; 2 specimens AM J20081, 35°02°S 151°06’E to 34°58’S 151°08’E, off Shoalhaven, N.S.W., 439-420m; 2 specimens, AM J20082, 34°28’S, 151°19’E, N.S.W., 412m; 1 specimen, AM J7155, off Eden, N.S.W.; 1 specimen, AM J12871, 35°44’S, 150°37’E, east of Brush Island, NES eeooamesle specimen, AM Ji3202 38°12" 5 149949 Bi to 38°ll7S: 14925375, southeast of Gabo Island, Vic., 439m; 1 specimen, AM J7007, off Newcastle, N.S.W.; 1 specimen, AM J20083, 36 miles south of Mt Cann, Vic., 205m; 1 specimen, AM J7052, off Eden, N.S.W. Distribution: Southeastern Australia, from off Newcastle, New South Wales to Bass Strait and the Great Australian Bight, in depths ranging from 128-457m. Etymology: Named for the Federal Fisheries Investigation ship ‘Endeavour from which extensive collections of southeastern and southern Australian fauna, including echino- derms, were made. Description: The holotype has 5 rays, inflated proximally and tapering to a fairly acute tip, R=40mm, r=7mm, R/r=5.7. The disc is small. The madreporite is small and ridged, occurring interradially at about %r. The abactinal skeleton (Figs 2A, 4A) is stout and closely reticulated, with 1-2, rarely 3 papulae per mesh. The overlapping plates are irregular — ovoid to stellate in shape. Additional small plates and processes occasionally project from the main skeletal plates into the papular region. The plates are raised centrally into low ridges bearing spinelets in tufts of 3-9, or in 2-3 irregular crescentic rows (Fig. 4B). The spinelets are fine, serrated and taper to an acute tip (length: 0.24mm, width: 0.06mm). PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 86 NEW ECHINASTERID ASTEROIDS Fig. 2. Odontohenricia endeavourn n. gen. et. sp. (holotype; AM J8792): A. abactinal surface; B. actinal surface (R=40mm). PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 87 F. W. E. ROWE AND E. L. ALBERTSON yy 4 u SC j i) ] region ina Beach farm 1ew O AM J8792); A. lateral vi d papulae. ? . gen. et. sp. (holotype , adambulacral sp wn icla endeavourt 3. Odontohenr 18. show ines an ines ] oral sp ing apica PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 88 NEW ECHINASTERID ASTEROIDS B proximal D> proximal P C proximal DP Imm 1Pi8!p Fig. 4. Odontohenricia endeavour n. gen. et. sp. (holotype; AM J8792): A. denuded abactinal skeleton of arm, (proximal); B. abactinal spinelets of arm (proximal); C. denuded skeleton of arm, lateral view from 4R, dis- tally; AB = abactinal plates, S = superomarginal plates, IT = intermarginal plates, IF = inferomarginal plates, AC = actinal plates, AD = adambulacral plates; D. profile view of 2nd adambulacral plate showing alignment of furrow spines and subambulacral spines and with adjacent actinal plates; E. actinal view of pair of oral plates showing large, apical, recurved oral spine and suboral spines. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 FW. E. ROWE AND E. L. ALBERTSON 89 The marginal plates (Figs 3A, 4C) are distinct due to their size, regular shape and alignment. The superomarginals are quadrilobed, about as long as wide, with a median spinelet bearing ridge oriented obliquely. The inferomarginals are also ridged and quadrilobed, markedly higher than wide, making this series conspicuous. The intermarginal plates are more or less quadrilobed, bridged with smaller plates. Both the quadrilobed and bridging plates are ridged centrally and bear spinelets. The principal intermarginal series extends %R, while a second series consists of a few plates only. Actinal plates occur in two rows, the first extending almost the ray length, the second only %4R. The plates are centrally ridged and bear spinelets. There is one papula present between adjacent plates to the marginal series. A vertical row of 2-3 tapered blunt furrow spines is present within the furrow. The adambulacral plates (Figs 3B, 4D) bear 2 transverse rows of up to 5 cylindrical sub- ambulacral spines on their actinal surface. The spines decrease in size across the plate. Spines at furrow edge are cylindrical, rounded apically, while those behind are tapered, serrated becoming similar to spinelets on adjacent actinal plates. Each pair of oral plates is dominated by a large recurved hyaline spine at their common apex, 4(5) dentate spinelets occur on the furrow margin of each plate with 3-4 suboral spines on the actinal surface of each oral plate. Paratypes: Some variation was found within the 11 designated paratypes. All were 5 rayed R=30-69mm, r=5-12mm except one (AM J3072) which is 6 rayed. The number of intermarginal and actinal series varied from 1-3, extending from %-%R, in the first series, a few plates to %R in the second, and a few plates sometimes present as a third. Similarly, the first actinal series may extend between %R to R, the second from a few plates to %R, and sometimes a few plates formed a third series. Other material: Some 15 further specimens have been identified. These fall within the variation we recognize within the type series. Remarks: O. endeavouri differs from other known Odontohenricia by the arrangement of the abactinal spinelets, the presence of stellate plates in the abactinal skeleton and the elongated inferomarginal plates. Odontohenricia clarkae n. sp. Figs 5A-B, 6A-E Diagnosis: Rays 5, R=26-33mm, r=4.4-6.6mm, R/r=5-6; rays cylindrical, tapering to a rounded tip; abactinal skeleton reticulate, 2-9 papulae per papular area; plates ridged medially, carrying 6-15 serrated spinelets; marginal plates distinct, with obvious vertical and horizontal alignment of series; plates quadrilobed, with trilobed ridges; infero- marginal plates higher than broad, more elongate than superomarginals, 1-2 intermar- ginal series; 2-4 actinal series; 1-5 papulae present throughout marginal series, adambulacral plates with 1-2 furrow spines in vertical series, one prominent sub- ambulacral spine on adradial edge of plate with 10-13 subambulacral spines behind in 2 rows; oral plate pair with large apical recurved spine, each plate with 2-3 small spinelets along furrow margin, and an additional 3 suboral spines. Material examined: Two specimens, holotype and paratype, BM (NH) 1951.5.8.2, 34°33’S 18°20’E, off the Cape of Good Hope, South Africa, 290m. PROG. LINN. SOC. N.S.W., 110 (1), (1987) 1988 90 NEW ECHINASTERID ASTEROIDS tf ‘th sm Fig. 5. Odontohenricia clarkae n. gen. et. sp. (paratype; BM (NH) 1951. 5.8.2): A. abactinal view; B. actinal surface, showing apical oral spines, abactinal and marginal spinelets (R= 26mm). PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 F. W. E. ROWE AND E. L. ALBERTSON Oi B distal [> ) . distal [> <]distal Imm distal [> Fig. 6. Odontohenricia clarkae n. gen. et. sp. (holotype; BM (NH) 1951. 5.8.2): A. denuded abactinal skeleton of arm (proximal); B. abactinal spinelets of arm (proximal); C. denuded skeleton of arm, lateral view (proximal); AB = abactinal plates, S = superomarginal plates, IT = intermarginal plates, IF = infero- marginal plates, AC = actinal plates, AD = adambulacral plates; D. profile view of 5th adambulacral plate with furrow and subambulacral spines; E. oblique-lateral view of pair of oral plates showing large apical oral spine and suboral spines (R=33mm). PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 92 NEW ECHINASTERID ASTEROIDS Distribution: At present known only from the type locality. Etymology: Named for Ailsa Clark for her significant contribution to the knowledge of the echinoderm faunas, including that of southern Africa. Description: The species has 5 rays, cylindrical and tapering to a rounded tip. Holotype R=33mm, r=6.6mm, R/r=5; paratype R=26mm, r=4.4mm, R/r=5.9. The disc is small. The madreporite occurs interradially at approximately r. The abactinal skeleton (Fig. 6A) is delicate and reticulate, with 2-9 papulae per papular area, but more commonly 5. The plates are irregular, ranging from bar to crescent shaped, the latter contributing to a scalloped appearance of the abactinal sur- face (Fig. 5A). Each plate is raised centrally into a low ridge, bearing 6-15 spaced, fine, serrate spinelets (0.26mm long, 0.04mm wide) (Fig. 6B). The marginal plates (Fig. 6C) are distinct. The plates and their medial ridges are regular in shape and alignment forming a well developed horizontal series. The supero- marginals are quadrilobed, higher than broad and each bears a central crescentic to tri- lobed ridge which carries 6-16 spinelets. The inferomarginals are quadrilobed, being markedly higher than broad. Spine-bearing medial ridges occur similarly to those on the superomarginals. The intermarginal plates are irregularly lobed, bearing vertical or crescentic, spine-bearing, medial ridges, the series extending °/5R in the holotype, and in the paratype 2/5-’%4R. The holotype has 2 series of intermarginal plates, the second extending to %R. The actinal plates are irregular and quadrilobed, with spined median ridges. The first actinal series extends for */5R and a second series, of a few plates in the holotype, and extending 4R in the paratype, lies above. The paratype also bears a series of a few plates between the second actinal series and the inferomarginal plates. Papulae are found throughout the marginal series (2-5 per papular area). Between the plates of the actinal series a single papula may be present. The spinelets on all marginal plates are similar to those of the abactinal plates, being irregularly toothed apically. The spinelets (12-18) are spaced on the ridge surface available (Fig. 6B). Within the furrow there is a vertical series of 1-2 smooth tapered furrow spines in the holotype but only a single spine occurs in the paratype. The adambulacral plates (Fig. 6D) bear one long smooth subambulacral spine at the furrow edge. Behind these stand 10-13 subambulacral spines, decreasing in height with distance from furrow. The smallest spinelets are similar to those on the abactinal and marginal plates. Each pair of oral plates (Figs 5B, 6E) is dominated by a large smooth recurved spine apically. In the paratype each plate also carries 3 spines along the furrow margin and 3 suboral spines on the actinal surface. Some of these spines have been lost in the paratype and completely lost from the holotype through mechanical damage. A small smooth spine is present distally on the lateral face of the oral plate. Remarks: The variation seen between the holotype and paratype can be attributed to size differences. The crescentic plates abactinally give a scalloped effect not seen in other species of Odontohenricia here described. This, with the combined features of the trilobed ridges of the superomarginal and inferomarginal plates, the marked vertical alignment of the marginal series and oral plate armature distinguish this species. The two specimens described herein were among 3 identified as Henricia abyssalis’ (Perrier), a species known to occur in South African waters (A. M. Clark and Courtman-Stock, 1976). PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 F. W. E. ROWEANDE. L. ALBERTSON 93 Odontohenricia fishert n.sp. Figs 7A-B, 8A-E Henricia leviuscula annectens; Fisher 1911, 291 (part) (non H. leviuscula annectens Fisher 1910). Diagnosis: Rays 5, R=82mm, r=15mm, R/r=5.5, inflated proximally, tapering distally to narrow tip; abactinal skeleton open reticulate, plates ridged medially, 1-5 accessory plates sometimes present in papular areas, 1-7 papulae per area; abactinal spinelets 0.4mm long, 0.08mm wide, tapering to serrated tip, spinelets densely clustered; margi- nal plates distinct, quadrilobed, ridged medially, spinelets similar to those on abactinal plates; inferomarginals slighty higher than superomarginals; 4 rows of intermarginal plates, principal row extending for ‘2R; 3 rows of actinal plates extending */5R, !/5R and !/10R respectively; papulae extend to actinal series, with 1 per area; 2 furrow spines in vertical series; adambulacral plate with 3-5 smooth, cylindrical, sometimes spatulate subambulacral spines occurring on the adradial edge, behind which occur 9-11 cylindri- cal spines, apically serrate or smooth, in 2-3 more or less distinct rows, these sub- ambulacral spines decrease in size across the plate; a large recurved spine occurs at the apex of the jaw, with 12-15 smaller, smooth suboral spines occurring on the adradial and actinal surfaces of the oral plates. Material examined: One specimen, holotype, U.S. National Museum E3838, 48°33’N, 124°53’W, off Washington, U.S.A., 108m. Distribution: At present known only from the type location. Etymology: Named for Dr W. K. Fisher who has contributed much to our knowledge of the classification of sea-stars. Description: Rays 5, long and cylindrical, inflated proximally and tapering to rounded tip (Fig. 7). R=82mm, r=15mm, R/r=5.5. The disc is relatively small. The madreporite, is ridged and spine-bearing, occurring interradially, at about Yr. The abactinal plates are irregularly bar to crescent shaped and form an irregular reticulum. Frequently (at least proximally) additional small plates are present within the papular areas (Fig. 8A). These areas may contain 1-7 papulae, but more commonly 3-4. The skeletal plates are raised medially forming low ridges. These ridges bear dense clusters of 3-24, but more commonly 15-20 tapered spinelets (length: 0.4mm, width 0.08mm) which are serrated apically (Fig. 8B). The marginal series are distinct, the plates being aligned vertically and horizon- tally (Fig. 8C). The superomarginal plates are quadrilobed, higher than broad, each with an oblique crescentic ridge bearing a dense cluster of spinelets (24-28). The infero- marginal plates are quadrilobed, as high as wide, but larger than the superomarginals. Medial ridges of these plates are crescentic to ovoid in shape and carry dense clusters of spinelets (23-34). The intermarginal plates are somewhat rectangular but may be enlarged basally or lobed laterally, and bear circular to ovoid medial ridges bearing spinelets. One inter- marginal series extends approximately %R although not clearly defined throughout. A second series between the first intermarginal and the inferomarginal series extends '/e6R. In addition, a few plates occur between the superomarginal series and the prin- cipal intermarginal series, and also between the second intermarginal and inferomargi- nal series. The actinal plates are quadrilobed, as high as wide, each with an ovoid, medial, spinelet-bearing ridge (13-26 spinelets). The first series extends */5R, a second PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 94 NEW ECHINASTERID ASTEROIDS / ni y Wy, ‘uff Fig. 7. Odontohenricia fisheri n. gen. et. sp. (holotype; U.S. National Museum E3838): A. abactinal surface; B.actinal surface, showing apical oral spines, actinal papulae and adambulacral and actinal spinature (R=82mm). PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 F. W. E. ROWE AND E. L. ALBERTSON 95 .?: Proximal P \. QN, Proximal > J distal Fig. 8. Odontohenricia fishert n. gen. et. sp. (holotype; U.S. National Museum E3838): A. denuded abactinal skeleton of arm (proximal); B. abactinal spinelets of arm (proximal); C. denuded skeleton of arm, lateral view from 11th to 16th superomarginal plate; AB = abactinal plates, S = superomarginal plates, IT = inter- marginal plates, IF = inferomarginal plates, AC = actinal plates, AD = adambulacral plate; D. profile view of 9th adambulacral plate showing furrow and subambulacral spines and with adjacent actinal plate and spinelets; E. oblique-lateral view of oral plate pair with large apical and suboral spines. PROG. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 96 NEW ECHINASTERID ASTEROIDS series to !/5R, and a third to !/10R. The spinelets occurring on the marginal plates are similar to those found abactinally. Papulae extend throughout the marginal series with one papula between actinal and inferomarginal plates and elsewhere 1-5 per papular area. There are 2 spines in vertical series within the furrow. The adambulacral plates bear 3-5 smooth, elongate and sometimes spatulate spines. Behind these are 9-11 cylin- drical spines which may be serrated apically or smooth. These form 2-3 more or less distinct rows, although they may also be clustered (Fig. 8D). Each pair of oral plates bears a large, smooth, recurved, hyaline spine apically (Fig. 8E). Each oral plate also bears 5-6 small cylindrical spines along the furrow margin, with an irregular row of 4-6 subambulacral spines on the actinal surface of the plate. In addition there are 3-5 subambulacral spines distally. Remarks: Odontohenricia fisheri is clearly distinguishable from other species of Odonto- henricia by its size, the presence of additional plates within the abactinal reticulum, the rectangular intermarginal plates and the abundance of oral plate spines and other skele- tal plate spines. Fisher (1911) identified this specimen as Henricza leviuscula annectens. Odontohenricia hayashi n.sp. Figs 9A-B, 10A-F Diagnosis: Rays 5, R=35mm, r=6mm, R/r=5.9mm; slender, tapering to rounded tip; abactinal skeleton reticulate, 1-5 papulae per area; abactinal spinelets, 0.48mm long, 0.08mm wide, slender and apically dentate, 1-9 spinelets per plate in tufts or irregular rows; marginal plates prominent, superomarginal, inferomarginal and actinal series being similar in shape size and alignment, all with oblique medial ridges bearing spine- lets; intermarginal plates irregular, lobed, frequently fused, in 3 series, first row to 4R and the second and third comprising a few plates only, 2 actinal series to 4R and !/5R respectively, papulae present between marginals and actinals, 1-4 per papular area; plates show widespread extensions which restrict the papular area, marginal spinelets as for abactinal plates; 1(2) furrow spines, adambulacral plates with a single prominent subambulacral spine on adradial edge, and with 8-10 cylindrical subambulacral spines in, more or less, 2 rows, decreasing in size across the adambulacral plate; oral plates with recurved, hyaline, apical spine, additional 7(8) regular suboral spines on each plate. Material examined: one specimen, holotype, U.S. National Museum E34052, Sagami Bay, Honshu, 35°50’N, 139°35’E, Japan, 152-289m. Distribution: at present known only from type locality. Etymology: named for Dr Ryoji Hayashi, who has contributed much to our knowledge of the Japanese sea-stars. Description: Rays 5, cylindrical and slender, tapering to a rounded tip (Fig. 9). R=35mm, r=6mm, R/r=5.9mm. The disc is relatively small. The madreporite occurs interradially, %R. The abactinal plates are tapered, bar and quadrilobed in shape, overlapping to form a fine, open, more or less regular reticulate skeleton (Figs 10A, B). The plates are raised medially and bear 1-9 but more usually 5-7 spinelets in tufts or irregular rows. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 F.W. E. ROWE AND E. L. ALBERTSON Fig. 9. Odontohenricia hayashi n. gen. et. sp. (holotype; U.S. National Museum 38307): A. abactinal surface; B. lateral view of arm (R=35mm). PROC. LINN. SOC. N.S.W,, 110 (1), (1987) 1988 98 NEW ECHINASTERID ASTEROIDS proximal > Proximal p> Proximal > C E F Fig. 10. Odontohenricia hayashii n. gen. et. sp. (holotype; U.S. National Museum 38307): A. denuded abactinal skeleton of arm (proximal); B. denuded abactinal skeleton of arm (distal); C. abactinal spinelets of arm (proximal); D. denuded skeleton of arm, lateral view from 7th to 13th superomarginal plate; AB = abactinal plates, S = superomarginal plates, If = intermarginal plates, IF = inferomarginal plates, AC = actinal plates, AD = adambulacral plate; E. profile view of adambulacral plate showing furrow and subambulacral spines and with adjacent actinal plate (composite drawing from 13th and 14th plates due to damage); F. oblique-lateral] view of oral plate pair with large apical spine and suboral spines. PROC. LINN. SOC. N.S.W., 110 (1),.(1987) 1988 FW. E. ROWE ANDE. L. ALBERTSON 99 The spinelets are 0.48mm long, 0.08mm wide and dentate apically. The papular areas are small, bearing 1-5 but more commonly 2-3 papulae per area. The superomarginal, inferomarginal and actinal plates are distinct, being of comparable shape and size, and regularly aligned (Fig. 10D). They are quadrilobed, as high as wide, and bear a transverse, medial, spinelet bearing ridge. The intermarginal plates are irregularly lobed and frequently fused. There is one principal series, extend- ing %R, and 2 additional series of only a few plates, one above and one below the prin- cipal series. These plates are also spined and ridged medially, and are aligned with those of the other marginal series. There are 2 actinal series of plates, the first extending %4R and a second extending '/5R. Papulae are present between these marginal series, usually 1 per papula area, rarely 2. There is extensive broadening of plates throughout the marginal series, excluding spaces for papulae. The spinelets found on the marginal plate ridges are as those of abactinal plates. Within the furrow there is one, sometimes 2 cylindrical spines in vertical series. There is a single prominent, cylindrical sub- ambulacral spine on the adradial edge of the adambulacral plate, behind which are 8-10 cylindrical, round-tipped subambulacral spines in 2 more or less regular rows (Fig. 10E). These spines decrease in height and breadth with distance from the furrow. The oral plate pair is dominated by a large smooth recurved apical spine. Four regular spines are also found along the furrow margin of the oral plate, with 1-2 additional suboral spines (Fig. 10F). Remarks: This species is clearly distinguished from other Odontohenricia here described by the small clusters of long abactinal spinelets, and the extension to the marginal plates. A. H. Clark identified 2 specimens from ‘Albatross’ station 3749 as Henricia densis- pina (Sladen) one of which we describe here as the holotype of O. hayashit. The identity of the second specimen may also be questionable though H. densispina has been recorded from Japanese waters by Hayashi (1940). ACKNOWLEDGEMENTS We wish to thank the following colleagues for the loan of specimens. Miss A. M. Clark, British Museum (Natural History), London, U.K.; Dr D. L. Pawson and Miss M. Downey, United States National Museum (Smithsonian Institution), Washington, D.C., U.S.A.; Ms Sue Boyd, Museum of Victoria, Melbourne, Vic., Australia. We also thank Mr Ken Graham for collecting material on FRV ‘Kapala and New South Wales State Fisheries for donating the specimens to the Australian Museum. The senior author wishes to acknowledge support from the Marine Sciences and ‘Technologies Grant Scheme (MST: 84/2092). References CLaRK, A. M., and COURTMAN-STOCK, J., 1976. — The Echinoderms of Southern Africa. London: British Museum (Natural History). 1-277, 276 figs. CLARK, H. L., 1916. — Report on the sea-lilies, star fishes, brittle-stars and sea-urchins obtained by the F.LS. ‘Endeavour’ on the coasts of Queensland, N.S.W., Tasmania, Victoria, S. Australia and W. Australia. Endeavour Res. 4:1-123, 11 figs, 44 pls. ——, 1946. — The echinoderm fauna of Australia. Publs. Carnegie Instn No. 566: 1-567. FISHER, W. K., 1910. — New starfishes from the North Pacific. II. Spinulosa. Zool. Anz. 35: 568-574. ——,, 1911. — Asteroidea of the North Pacific and adjacent waters Part 1. Phanerozonia and Spinulosa. Bull. U.S. natn. Mus. 76: 419pp, 122 pls. ——, 1940. — Asteroidea. Discovery Rep., 20: 69-306, 23 pls. Gray, J. E., 1840. — A synopsis of the genera and species of the Class Hypostoma (Astertas Linn.). Ann. Mag. nat. Hast. (1) 6: 175-184. PROG. LINN. SOG. N.S.W., 110 (1), (1987) 1988 100 NEW ECHINASTERID ASTEROIDS HayasHI, R., 1940. — Contributions to the classification of the sea-stars of Japan. I. Spinulosa. J. Fac. Scz. Hokkaido Imp. Univ. (6) 7(3): 107-204, pls vii-xi1. PERRIER, E., 1891. — Echinodermes. I. Stellerides. Rept. Mission sct. du Cap Horn 1882-83. 6 (3) Zoologie: K1- K198, pls. 1-13. SLADEN, W. P., 1889. — Asteroidea. Rep. scient. Results Voy. ‘Challenger’ (Zool.). 30: 893pp, 117 pls. Rowe, F. W. E., and ALBERTSON, E. L., 1987a. — The echinoderm genus Henricia Gray, 1840 (Asteroidea: Echinasteridae) in southern and southeastern Australian waters, with the description of a new species. Proc. Linn. Soc. N.S.W. 109: 183-194. , and , 1987b. — A new species in the echinasterid genus Echinaster Muller and Troschel, 1840 (Echinodermata: Asteroidea) from southeastern Australia and Norfolk Island. Proc. Linn. Soc. N.S.W. 109: 195-202. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 ERRATUM HELENE A. MARTIN Presidential Address. Cainozoic History of the Vegetation and Climate of the Lachlan River Region, New South Wales. Proc. Linn. Soc. N.S.W. 109 (4), (1986) 1987: 213-257. p.244, lines 45-46, should be: Today, the mean annual precipitation for Hillston is about 350mm, Forbes 520mm to 535mm and Cowra 610mm to 630mm. PROC. LINN. SOC. N.S.W., 110 (1), (1987) 1988 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 110 NUMBER 2 als teh 3 Ea a any Wert PEE, SPR WILLIAM MAGCLEAY MEMORIAL LECTURE 1987 The elastic-sided Gumleaf, or: The Rubber Cuticle and other Studies of the Corymbosae STELLA G. M. CARR and D. J. CARR Research School of Biological Sciences, Australian National University, Canberra [Delivered by D. J. CARR 30 September 1987] Introduction: the rubber cuticle of the Corymbosae In 1908, the Sydney phytochemist Henry G. Smith, of the Technological Museum, published an account of his analysis of the cuticles of Eucalyptus gummifera and those of two species of Angophora. He showed that, using suitable solvents, a material could be extracted from young leaves of these species which had chemical and physical properties identical with those of natural rubber (caoutchouc). It was soluble in chloroform, and in absolute ether but not in petroleum ether. Various tests, including melting point and ability to be vulcanized, convinced Smith that the material was indeed rubber. It has long been known that the cuticles of young leaves of species like E. gummuifera, spotted gum (E. maculata), and red flowering gum (E£. ficzfolia), are easily removed almost intact from the leaf surfaces and have the elastic properties of rubber. Later on, as the leaves mature and become fully developed, the cuticle undergoes changes which prevent it from being stripped off the leaves. This extraordinary discovery has not rated mention in any of the botanical textbooks, in the many reviews of recent decades on the topic of the plant cuticle or in two monographs devoted to that subject (Martin and Juniper, 1970; Cutler et al., 1982). The only reference we have been able to find is in the 2nd Edition of Metcalfe and Chalk (an expensive, poor relation of the magnificent first edition), in which Metcalfe (1983) reported that ‘the secretion of a rubbery substance from epidermal papillae on young shoots of three species of Angophora and fourteen species of Eucalyptus belonging to the section Corymbosae (Myrtaceae) has been reported by Welch (1923). Metcalfe thus added confusion to the already confused views of Welch, who 1n describing the papillate epidermis of the Corymbosae as a secretory tissue, ignored the fact that all shoot epidermes which secrete a cuticular layer are also, ipso facto, secre- tory tissues, whether papillate or not. Welch was somewhat misled by the fact that during embedding in paraffin the rubber tends to dissolve in the embedding medium and be at least partially lost. Metcalfe failed to follow up the reference to Smith’s work given by Welch. Moreover one of Welch’s 14 eucalypts is ‘E. santalzfolia’, which, as he points out, does not belong to the Corymbosae. Nor, as Blake (1953) later pointed out, should E. tessellaris, another of Welch’s species, be included in the Corymbosae. Welch did, at least, recognize the material covering the leaves as a cuticle and not, to use Metcalfe’s phrase, mere ‘rubbery secretions’ In his article on ‘Secretory Structures: cells, cavities and canals, Metcalfe appears to have followed Nelsonian principles in turning a blind eye to much of the recent literature. In 1956, when we commenced our studies of the biology of the eucalypts, we repeated Smith’s extractions and most of his tests, using young developing leaves of E. ficifolia grown in cultivation in Melbourne. We were not only able to verify Smith’s PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 102 THE ELASTIC-SIDED GUMLEAF 100 = <<) L Hevea Sie (= ° < [ fo SH) E L w Cc oa! | IS L | Eucalyptus ficifolia 0 | 1 [ee TE 1 1 2 3 4 5 6 7 8 9 10 "1 12 13 14 15 ] Wavelength (um) 250 200 Mimusops palit (gutta) Lon S ~ im Sew 2 ; i i?) (SF e FE 150 100 ee 11.50 12.0 12.50 7) Wavelength (Um) Fig. 1. Infra-red transmission spectrograms; abscissa, microns. 1. Rubber from Hevea brasiliensis and a cuticle preparation from Eucalyptus ficifolia. 2. Comparisons between rubber from various sources, gutta percha, and a cuticle preparation from Eucalyptus ficifolia. (Data courtesy Dr T. P. O’Brien and the late Sterling B. Hendricks). claims, but we also were able to produce little balls of rubber with a very satisfactory bounce. The most favoured method for the identification of rubber is infra-red absorp- tion spectroscopy. Dr T. P. O’Brien, a former student of ours, then employed at ICI in Melbourne, ran some IR absorption spectra of whole cuticles and arranged to send them for comment to Sterling B. Hendricks at the Beltsville Laboratory of the US Department of Agriculture (Fig. 1). Dr Hendricks, by training a physicist and a spectro- PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G. M. AND D. J. CARR 103 scopist who had worked with Linus Pauling, had been involved during the last World War in a programme of investigation of various plants (e.g. guayule and the kok-saghyz dandelion) as sources of natural rubber. He examined the IR spectra and in reply to Dr O’Brien stated that ‘your samples appear to me on the basis of absorption in the region of 12 microns (a critical region) ‘to be rubber. More recently we repeated the extraction of rubber from young leaves of EF. calophylla sent to us by Mrs M. J. Hamersley from Western Australia. Using the samples thus obtained we carried out the chemical test known as Weber’s test for rubber. A small sample, 50mg, is cut into fine pieces and put into a test tube. A drop of bromine is added and the test tube is warmed for 30 seconds on a water bath. Then a gram of solid phenol 1s added. Natural rubber gives a violet coloration, but synthetic rubbers, such as neoprene, give only a weak reaction. Needless to say, the rubber samples from E. calophylla gave a strong positive reaction. The two eucalypt species mentioned so far belong to a group commonly called bloodwoods, or botanically the Corymbosae. Following Smith’s publication, Maiden (1908-1928) and Welch (1923) claimed to have observed rubber in a variety of species of eucalypts in other taxonomic groups. We have no evidence on this point, although it is inherently not unlikely. The rubber cuticle is layered In a number of ways, the cuticle of the Corymbosae is different from that of other angiosperms. The consensus of opinion appears to be that, in angiosperms generally, the cuticle steadily increases in thickness as the leaf or stem grows. In the Corymbosae the cuticle of the leaf increases in thickness only until the leaf 1s between a third and a half fully-grown. It then decreases in thickness until near maturity (Fig. 2). The parameters on which these graphs were constructed are discussed in our recent book, Eucalyptus IT. We would emphasize that much more information ( data on plastochrone duration, measurements of the duration and extent of lamina growth, measurements of average cuticle thickness during growth) is needed to establish and quantify these concepts but there can be no doubt that the cuticle thickness does increase and then decrease. This can be seen from a hand-section of a leafy bud of a species such as E. gummifera. Clearly then, the rubber cuticle is laid down at an early stage on the surface of the leaf primor- dium and is subsequently stretched and thins out during what used to be called the ‘grand phase of growth’ of the leaf. The cuticle remains thick over the midrib and margins of the leaf which expand mainly in length and thickness, especially on the lower surface, (thus increasing its area relative to the upper surface) during the later phases of leaf growth. Welch measured the thickness of the cuticle on a leaf of E. gummifera less than 1 millimetre wide to be between 170 and 185u4m. Our own measurements of E. maculata grown in Canberra show a much more modest maximum thickness. Since the leaves appear in pairs in the buds, with the inner (adaxial) faces of each pair adherent, the thickness of the cuticle on those surfaces lags behind that of the freer abaxial sur- faces. Later on, the thickness of the cuticle on the upper surface catches up and at maturity is thicker than that of the lower surface. This is due to the fact that, at a late stage in leaf expansion, the epidermal cells begin to lay down layers of cutinized cuticle (Fig. 3). These develop to a greater thickness on the upper surface than on the lower. They are also thicker on the midrib and margin than elsewhere. Stomata are initiated and begin to break through to the outer surface of the cuticle at or just before the time the leaf is half fully-grown, while the cuticle is beginning to be stretched and thin out. All the stomata are already formed by the time the cuticularized layers begin to be laid down. We shall return to the stomata later on. The flower bud and its constituent parts, such as the sepals, petals, style, stamens and the loculi of the ovary are also covered with PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 104 THE ELASTIC-SIDED GUMLEAF thickness (um) abaxial rubber cuticle G 0 10 20 30 40 50 60 time (days) 100 ac ss 5s 80 1S) 2 Oo = 00 g ss) Ze AO = St © 20 G 0 3 0 10 20 30 40 50 60 time (days) Fig. 2. Graphs to show changes in the thickness of the abaxial (upper graph) and adaxial cuticle of an adult leaf of E. maculata. The lines on the lower right of each graph beginning after c.32 days, show the formation of the cuticularized layers underlying the rubber cuticle. a rubber cuticle, which is often massively thick, and, at maturity, is not underlain by cuticularized layers. By suitable staining one can demonstrate that the whole of the thick rubber cuticle consists of layers, each consisting of a thin dark layer and a thicker lightly-staining one (Fig. 4). It is often possible to count the layers in a section of the leaf and in a leaf of E. maculata estimated to be between 30 and 40 days old, when the cuticle of the adaxial surface is at its maximum thickness, the number of layers is between 30 and 40. ‘This is PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 105 Fig. 3. Diagram to show zonation in the cuticle of a mature, fully-grown leaf of a species of Corymbosae. 1, cell wall. 2, 3 cuticularized layer, layer 3 with radial striae. 4, 5 zones of rubber cuticle, the layers well-spaced in 4, closely-spaced in 5. 6, the ‘cuticle proper’. at least presumptive evidence that each bilayer is the product of a single day’s secretory activity on the part of the epidermis. A similar diurnal layering is well-known for the cell-walls of certain algae and for the hairs on the seeds of cotton. Moreover, the cuticles of insects have diurnally-produced layers (Neville, 1963). The difference in density between the two portions of a bilayer may reflect different packing densities of the rub- ber in them or perhaps differences in ancillary materials associated with the layers. Again, we wish to stress that much more work is needed to establish the diurnal rhythm in rubber production, the nature of the differences in the bilayers, the nature of ancillary materials in the rubber cuticle etc. Stomatal breakthrough in the Corymbosae As we have already shown in studies of the stomatal development in other groups of eucalypts, the stomatal initials appear and the full structure of the guard cells develops, including the split between them, while the guard cells are completely covered by an un- broken cuticle (Fig. 5). Above the line of closure of the guard cells a split must develop in the cuticle to give access to the atmosphere. We have provided evidence from light microscopy which supports the view that this split, resulting in stomatal breakthrough, develops by a process of digestion of the cuticle above the guard cells, probably by enzymes secreted by them into the cuticle (Carr and Carr, 1978; Carr and Carr, in preparation). The hole in the outer surface of the cuticle which leads to the guard cells we termed the ostzole (Carr and Carr, 1978). Although an essential part of the functional stoma, and, in thick cuticles, often smaller in area than the pore between the guard cells of the open stoma, it had not previously been given a name. Stace (1965) ignores it; Wilkinson (1979) calls it ‘the outer stomatal ledge aperture’, a long-winded appellation PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 106 THE ELASTIC-SIDED GUMLEAF Wy Py , iL yy, Yi) yh Fig. 4. Lamellate rubber cuticle of developing leaves of E. maculata. 1, 3, 4 adaxial surface, 2 abaxial surface. 2 leaf about half fully expanded; 3, 4 fully expanded. In 2, the number of bilayers (a light zone plus a dark line) is between 30 and 40. In 3 the asterisk indicates a widening of one of the light zones. In 4 arrows indicate positions where losses of outer layers can be observed. cp, cuticle proper; cl, cuticularized layer just beginning to be formed; w, cell wall. Scale bars, 10um. which is wrong for sunken stomata (e.g. those of Proteaceae spp., Aloe spp., Ficus spp., Eucalyptus incrassata (Carr and Carr, 1978), species of the Brsectae (Eucalyptus) and of the Lehmanmanae (Eucalyptus) (Carr and Carr, 1980c), which have outer stomatal ledges which are not fused with the cuticle. The correct term for the aperture between the outer stomatal ledges is the ezsodial aperture. The problems of stomatal breakthrough in the PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 107 Fig. 5. Stomatal breakthrough, giant stomata. Developing leaf of E. maculata, abaxial surface. In 2 the guard mother cell (gmc) has divided, giving rise to the guard cell initials. Scale bar, 10um. Corymbosae are quite different from those we have observed in other eucalypts, or in common mesophytic plants such as Phaseolus and sunflower (Carr and Carr, in prepar- ation), and are solved in quite a different manner, because of the presence of a layered, rubber cuticle above the stomatal complexes. In the Corymbosae, the guard mother cell (GMC) expands and produces a vacuolate apical swelling or papilla which extends into the cuticle above it. As it touches the layers of rubber they appear to dissolve and their cut edges adhere to it (Fig. 5 [1]). The GMC divides anticlinally to yield the two guard cell initials (Fig. 5 [2]). Next the apical swell- ing relaxes so that the outer surface of the developing guard cells resumes a position level with the rest of the epidermal cells (Fig. 6 [1]). Still within the mother cell envelope, the two guard cells begin to lay down the layers of their upper and lower wall thickenings. These become cutinized (Fig. 6 [2]). They will eventually form the inner and (part of) the outer stomatal ledges. At this stage, above the developing guard cells a conical zone appears in the cuticle (Fig. 7). The zone is filled with a granular precipitate. It is evi- dently a region in which dissolution of the rubber layers takes place, the precipitate being, presumably, a product of the digestion of the rubber. A similar conical zone of dissolution has been reported by us as appearing above the developing guard cells of Eucalyptus incrassata, which has a thick cutinized (1.e. non-rubber) cuticle (Fig. 7 [2]). It is therefore timely to give such a zone a name; we have called it the conus, short for conus dis- solutzonts. Our interpretation of the origin of the conus is that the guard cells, still en- closed within their mother cell envelope, secrete into the cuticle enzymes capable of digesting the cuticle, whether it consists of rubber (as in the Corymbosae) or of cutinized layers. Very probably the GMC itself has some of the necessary enzyme (we may call it a PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 108 THE ELASTIC-SIDED GUMLEAF Fig. 6. As Fig. 5, to show later events in stomatal breakthrough. In 1 the guard cell initials have withdrawn dragging with them the innermost rubber lamellae. 2, upper and lower guard cell thickenings have begun to form inside the old gmc envelope. ‘laticase’ in the Corymbosae) on its outer surface and that this is the reason for its ability to break through and attach to itself the rubber layers it touches. As the conus enlarges and the number of still undigested rubber layers above it decreases, it begins to intersect with the outermost layers of rubber. At this stage, scan- ning electron microscopy of the leaf surface (Fig. 8 [1]) shows a number of black annuli (black because the thin cuticle is not as electron-reflective as elsewhere). Within such an annulus a circular line of breakage appears and the inner disc of rubber is released, shrivels and is lost (Fig. 8 [2]; Fig. 10 [4,5,6]). A more-or-less circular opening 1s left in the cuticle, leading down, through similar jagged holes in successive layers, to the guard cells. Inverted cuticles show these layers and the jagged edges of the holes made in them Fig. 8 [6]). This method of stomatal breakthrough applies to the earliest stomata formed on the leaf. These stomata, because they have an unusually high complement of sub- sidiary cells, are referred to as geant stomata. We shall have more to say about them in the second part of this lecture. Stomata formed later on, when the cuticle is rapidly thinning, also each have a conus but it becomes stretched out by the lateral expansion of the cells of the stomatal complex and is shallow (Fig. 9 [2]). Where it intersects with the surface layers of the cuticle it presents more the appearance of a small dot or a line (Fig. 11 [3,4,5]). Cracks appear along such a line and join up to form a slit. No disc of rubber 1s released. As leaf expansion continues, the slit widens to become an ellipse (Fig. 11 [8,9]), often still show- ing at its ends the traces of the line which initiated it. Inverted cuticles show the shallow cavities in the cuticle, above the ordinary stomata (Fig. 8 [5]). Breakthrough in these ordinary stomata brings the outer thickenings of the guard cells in contact with the outermost layers of rubber (Fig. 9 [4]) and they fuse with them to form a composite outer PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 109 Fig. 7. All except 2 E. maculata, to show the initiation of the conus dissolutionis (arrowheads) outside the gmc envelope. osl, outer stomatal ledge. 2. The conus (arrows) above a developing stoma of Eucalyptus incrassata. White arrowhead (2), layering of the cuticularized cuticle. Scale bars, 10p. PROC. LINN. SOG. N.S.W., 110 (2), (1987) 1988 110 THE ELASTIC-SIDED GUMLEAF y) Wh) Wy) Fig. 6. Stornatal breakthrough, scanning electronmicrographs E. erythrophloia. 1. Arrows, dark circles indicat- ing imminent breakthrough. 2. Detachment of the rubber disk, split appearing between two thickened rims of the annulus. 3. Breakthrough completed and peristomatal cuticular ornamentation beginning to form. 4. Through the ostiole, lamellae of the cuticle can be seen. 5 and 6. Detached, inverted cuticle taken from a leaf at a late stage in stoma formation. Cup-shaped hollows in the cuticle where stomatal breakthrough has oc- curred or is occurring. 6. One such area enlarged. Small arrowheads indicate lamellae in the cuticle. 0, osti- ole. Scale bars, all 10pm. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 111 ve Fig. 9. E. maculata, completion of stomatal breakthrough. 1. Large conus (arrow) of ‘giant’ stoma touching the outer surface of the cuticle. 2. Small, flattened conus of ‘ordinary’ stoma. Guard cells almost fully formed. 3. Completed ordinary stoma. Outer stomatal ledges formed as a composite of the guard cell thickenings and the outermost rubber layers. 4. ‘Ordinary’ stoma about to break through to surface. Asterisks in 2 and 4 indicate accumulation in anterior chamber of stoma of breakdown products of the dissolution of rubber lamellae. Scale bars, all 10um. stomatal ledge (Fig. 9 [3]). Occasionally there is a developmental hiatus, in which the final breakthrough does not take place (Fig. 11 [7]). The stomata, although otherwise fully- formed, are ‘blind’, i.e. have no opening or ostzole on the surface, and are therefore clearly non-functional. PROG. LINN. SOC. N.S.W., 110 (2), (1987) 1988 112 THE ELASTIC-SIDED GUMLEAF Fig. 10. Stomatal breakthrough, scanning electronmicrographs. E. calophylla. 1 and 2 dark rings, surrounding circular disks. 3. Arrows indicate splitting of the circular disk from the rest of the (wax-covered) surface. 4, 5. Arrows indicate shrivelled disks still attached to the ostiole or 6 loose on the surface of the cuticle. 7. Completed ostiole, through which one can observe (arrow) some of the lamellae of the cuticle. Scale bars, 1 10pm; all to same magnification. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 — S.G.M. AND D. J. CARR 113 Fig. 11. Stomatal breakthrough of ‘ordinary’ stomata. Scanning electronmicrographs. 1-8, E. calophylla, abaxial surface. 1 and 2. A field of stomata developing and breaking through. Note that the peristomatal cuticular ornamentation has begun to form (arrowheads). 3-6. Breakthrough beginning as a slit or crack (arrows) in the outer surface and 6 the ostiole completed. W, wax crystallites. 7. Light micrograph of leaf surface of a specimen of E. hamersleyana, showing (arrows) stomata with ostioles; the other stomata have no ostioles and are non-functional (‘blind’). 8. E. calophylla. After breakthrough, the ostiole enlarges and becomes elliptical. In 8 the outer stomatal ledges are visible through the ostiole. 9. E. eremaea. Fully-formed ‘ordinary’ stomata. Scale bars, all 10um. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 114 THE ELASTIC-SIDED GUMLEAF ui i By wy iy : < L. We ra Wi) Hj Wi] y GL LLL Fig. 12. Transverse sections of a leafy bud of E. collina. 1-5 transverse sections, 1 and 2 near the tip of the bud, 3 and 4 near its base. The leaf primordia are numbered in basipetal sequence, the precocious primordium of a pair with a prime (as e.g. 2’). b, b’, axillary buds (or, inb’, a position above an axillary bud), S, stem. Arrow- heads indicate positions where the cuticle of one primordium abuts on that of another. Scale bars, 1 1mm, all the others 10pm. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 » Pe ee ee ee a a a S.G.M. AND D. J. CARR 115 i 4 Me | # ty sa i A: i My | ! yy / j i yy aii fag 4 Fig. 13. E. collina; details of epidermal cells and cuticle lamellae. e, epidermal cell. In 1 arrows indicate abutment of cuticle of one primordium on that of another. Scale bars, all 100m. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 116 THE ELASTIC-SIDED GUMLEAF Fig. 14. E. collina, cuticle. 1 and 2. Tangential sections of cuticle showing concentric sections of domes of the cuticle over individual cells. Arrowheads show the envelope of a single cell, now divided to give two cells. 3-6. Transverse sections to show the adaxial cuticle (arrow in 1) of leaf primordia (numbered as before). The layers of cuticle are much more compact and consequently less domed than those of the abaxial surface. These less wavy layers are maintained as the outer layers of the adaxial surface on an older primordium 6. e, epidermal cell (its surface noticeably less ridged than those of the abaxial surface). Scale bars, 3 Imm, all the others 100pm. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 117 In our recently-published book, Eucalyptus IT, we have given a full account of these methods of stomatal breakthrough in the Corymbosae. We have also dealt with a third and quite different method, which is that followed by stomata initiated on the outer surface of the flower buds, as well as on the outer (abaxial) surface of the petals. Scurfiness of the cuticle Among the other consequences of the presence of a layered, rubber cuticle in the Corymbosae, 1s the possibility of scurfiness of the cuticular surface. Glaucousness in eucalypts is due to coverings of epicuticular wax in the form of tubes on the surface of leaves or flower buds. No species of the Corymbosae have tubular wax; those which have epicuticular wax have it in the form of wax platelets. It follows that none of the species of the Corymbosae can be glaucous. However the flower buds and, in a few cases, the young leaves of some species have a silvery sheen, usually described as scurfiness. The origin of scurfiness is explicable in terms of the layered cuticles of the Corymbosae. For its investigation, we have used a herbarium specimen of the tropical Western Australian species, E. collina. Fitzgerald, who discovered this species wrote of it that the branchlets and often the leaves appear ‘as if covered with frost’. “Uhe trees can be seen at a great distance, owing to their silvery whiteness, which is a distinct character of the plant. Dissections and transverse sections of leafy buds show that scurfiness is already apparent on the surface of the second smallest leaf primordia (P») and increases during the maturation of the leaves. After that, much of the scurfiness is gradually lost from the leaf surface, remaining, in the fully-developed leaves, mainly on the midrib and petiole. The innermost leaf primordia in the bud are tightly packed together forming what appears to be a solid mass, enclosed by the leaf cuticles (Fig. 12 [1]). For reasons we have already explained, and resulting from the close apposition of the adaxial surfaces of the innermost primordia, the abaxial cuticles become much thicker (at least initially) than the adaxial cuticles. The cuticle is strikingly layered, in much the same way as the cuticle of E. maculata (Fig. 13). On a P, leaf primordium, one can count 32 to 35 layers of the cuticle (Fig. 13 [4]). As in E. maculata, this suggests again a bilayer produced every day, assuming a plastochrone (interval between the production of successive leaf primordia) of about 8 days. Each layer consists of a series of domes, each produced over a single epidermal cell (Fig. 13 [1,2,5]). Each of these domes joins laterally with those of adjacent cells to form an undulating layer (Fig. 14 [4,5,6]). Cut tangentially (Fig. 14 [1,2]), the cuticle presents the appearance of a series of concentric circles, the domes of individual cells cut at various levels. Where a cell has divided, these circles are enclosed in ellipses represent- ing the now expanded and distorted domes of the original cell. Scurfiness appears as outermost layers begin to break down and exfoliate (Fig. 15). The outer layers appear to become too brittle to stretch to accommodate the continued expansion to the surface of the leaf primordium. Possibly the rubber oxidizes pre- maturely and loses elasticity. Cracks appear in the domes (Fig. 15 [2,3]) and the cracks spread, allowing rafts of detached cuticle to form (Fig. 15 [4]). Spaces between the detached cuticular layers fill with air, the source of the silvery sheen of the scurfy cuticle. Scurfiness of flower buds also arises from breakdown of the outer layers of rubber cuticle, as is discussed and illustrated in Eucalyptus II. Deep fissures develop in the cuticle and rafts of the outer layers may exfoliate. Discussion Following its initiation, the GMC ceases to form rubber. Indeed, its outer surface may be coated with a laticase. We had already shown for other eucalypts (not Corymbosae) PROG. LINN. SOC. N.S.W., 110 (2), (1987) 1988 118 THE ELASTIC-SIDED GUMLEAF Wy Me Hy Wy W : Wy / yl I an Fig. 15. E. collina. Development of scurfiness of the cuticle. 1. Location of early scurfiness (arrowheads), 2. Tangential section showing the erosion and breakup of the columns of the outermost cuticle lamellae. 3 and 4. Swelling of the light zones (arrowheads) and breakaway of the outer layers. Scale bars, 1 1mm, all the others 100pm. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. ANDD. J. CARR that, following its initiation, the GMC ceases to make cuticle above itself (Carr and Carr, 1978). Next, the guard cell initials which result from the division of the GMC begin to produce cuticularized layers within the mother cell envelope. The sequence of production first, of rubber layers, then of cuticularized layers of the cuticle is also that followed by the ordinary cells of the epidermis. This dual potentiality may have been a feature initially common to the ancestors of all the eucalypts but the capacity to produce rubber was reduced or lost in most groups of eucalypts. In some species perhaps only a few of the outermost layers of the cuticle consist of rubber (giving the cuticle a charac- teristic sheen) while the bulk of the cuticle consists of cuticularized layers. A rather similar situation 1s found developmentally in the leaves of certain deciduous species of the Corymbosae in the tropics. Following a few weeks of leaflessness, a new suite of thin, pale green leaves is produced which expand rapidly over a few days then gradually be- come thicker and darker green. In such leaves the initial cuticle of rubber is relatively thin and paucilamellate and, following the attainment of full size, a relatively thick cuticularized cuticle is developed to reinforce the protective layers. On this hypothesis, the rubber cuticle of the Corymbosae represents the survival of an ancient device for the protection of the leaf during its early expansion. This has given way, in most species of eucalypts, to the production of the less-overtly layered, cuticularized cuticles which are also a feature of most angiosperms. The flower is considered by morphologists to be a ‘conservative’ organ in relation to evolution. In the flowers of the Corymbosae only a rubber cuticle, which is in many instances remarkably thick and is always layered like that of the leaves, is formed. This supports the hypothesis that the rubber cuticle is, in an evolutionary sense, an older covering than the cuticularized layers. One can visualize that a rubber cuticle would present advantages to tropical species which, either deciduous or semi-deciduous, must rapidly expand a suite of leaves while still retaining a protective cuticle over their sur- faces. Unfortunately we know little about the cuticles of tropical plants, but it seems not unlikely that rubber cuticles may be found to be present in genera other than Eucalyptus and Angophora. A host of questions is raised by these findings. Are the bilayers really diurnally produced, and if so which of the two constituent layers is produced during the day and which during the night? Is layering suppressed in continuous light, as it is in insect cuticles and the cell walls of cotton hairs? Why are the cuticles of some species and some organs scurfy, those of other species and other organs not? Is the eventual hardening of the initially elastic cuticle due to oxidation, as was suggested by Smith (1908)? Perhaps scurfy cuticles lack antioxidants, such as a-tocopherol, a known constituent of eucalyptus oil; alternatively, since scurfy cuticles lack epicuticular wax, perhaps it 1s this lack which permits the development of scurfiness. Qualitative and quantitative aspects of the phytoglyph in Corymbosae In 1883 Ferdinand von Mueller wrote, concerning the plant fossils found in associa- tion with the deep leads of the Victorian goldfields, that: ‘By the aid of the microscope we may yet hope to be able to obtain characteristics of diagnostic value from the anatomy of leaves sufficiently positive to recognize ordinal and even perhaps generic groups. ‘How far this can be done even with living plants remains yet to be studied: but I was enabled, for instance, to demonstrate the occurrence of Epacridae in New Guinea from the microscopic comparison of the leaf epidermis, brought from thence without flowers or fruits, with the very peculiar cuticle of many Epacridae easily recognized microscopically’. In his Eucalyptographia Mueller illustrated features of the leaf epidermis of 39 species of eucalypts. He classified them as isogenous, with roughly equal numbers of PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 120 THE ELASTIC-SIDED GUMLEAF Fig. 16. E. nesophila and E. latifolia, showing different patterns of cuticular ornamentation. (Figs 16-23 all light micrographs of stained cuticles; in comparisons always the same surface, upper or lower of the two species, at the same magnification 1s shown.) Scale bar: 504m. iy yj 4, ri yi" Wy Wy) Wy 4), a i iy E. polycarpa upper Fig. 17. E. polycarpa, showing heterogenous arrangement of the stomata. Scale bar: 50um. See also legend to Fig. 16. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 121 stomata on both upper and lower surfaces, heterogenous, with many fewer on the upper than on the lower, or hypogenous, with stomata only on the lower surface. This excellent beginning of the study of the microscopic features of the leaves of Australian plants suffered a mortal blow when Maiden (1909-1928), after quoting Mueller’s stomatal classification, dismissed the study of stomata with the remark (our emphasis) that ‘the method cannot be used for diagnostic purposes because of the variation in the dis- tribution of stomata even in the same tree. Maiden offered no evidence of such variation and indeed made no further reference to stomata in his voluminous writings. Since that time no-one paid any attention to the features of the epidermis of eucalypt leaves until, in 1971, we published an account of our studies (Carr et al., 1971) of the leaves of the various ‘type materials’ supposed by Blake (1953) to be all of a single species, Eucalyptus dichromophiova. We were able to demonstrate that these materials belonged to no fewer than five different species. Blake had based his description of the juvenile foliage of EF. dichromophioia on leaves of a species of Angophora, included in the folders of one of these species of eucalypts. Recently, in our new book, Eucalyptus IT, we have shown that the type folders of E. erythrophloia (which included the Angophora leaves) also contain a speci- men of leaves of yet another species (E. ellipsordea), which is described for the first time. Thus the type folders of E. erythrophlova contain specimens, all collected in the same locality (where they still occur), of no fewer than three species belonging to two genera! The methods we used included scanning electron microscopy of the leaf cuticles, and light microscopy of stained cuticle preparations and of thin sections of cuticles. These approaches make it possible to reconstruct a 3-dimensional view of the cuticular patterns of the leaves and an analysis of the positions, shapes and types of cells in the leaf epidermis. For the totality of the information thus obtained we coined the term ‘phyto- glyph, which means literally ‘plant fingerprint. We maintain that, just as human finger- prints may be used diagnostically to identify persons, so the phytoglyph may be used to identify species of eucalypts, at least in some groups. Following the work on ‘E. dichromophloia, we used the method to identify old specimens, collected by Leonard Brass in Cape York Peninsula (and which had been filed under E. dichromophloia) as E. nesophila (S.G.M. Carr, 1972). At that time, this species was thought to be endemic to the Northern ‘Territory. Dr Nigel Wace then presented us with a problem in the form of a small leafy shoot collected on Fenelon Island, off the coast of South Australia. We identified the material as belonging either to E. socialis or to a closely related, possibly undescribed, species (Carr and Carr, 1976). At about the same time, that species, E. yalatensis, was described by C. D. Boomsma. Our further studies showed that the species of the informal group, the Brsectae, have several peculiarities of the leaf cuticle. In some species, during stomatal breakthrough, the cuticle becomes eroded into sinuous canyons or crypts, at the bottom of which lie groups of stomata (Carr and Carr, 1978, 1980b, 1980c). Other species of the Bisectae have special bars of cuticle forming the line of closure of the stomata (Carr and Carr, 1979). Yet others have peculiar ingrowths of cuticle in the anterior chambers of the stomata (Carr and Carr, 1980a). These features of the Bisectae were shown to be consistent with taxo- nomic relationships within that group. While such studies showed the usefulness of phytoglyphic methods, but by far the most successful and extensive use we have made of those methods has been in clearing up what M. R. Jacobs termed ‘the bloodwood puzzle’, i.e. the problem posed by the recognition and identification of the confusing and numerous tropical species of the Corymbosae. We have published accounts of these species in our two books, Eucalyptus I and Eucalyptus II. In doing so we refrained from adding to the volume of descriptive data by including all the data from the immensely detailed PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 122 THE ELASTIC-SIDED GUMLEAF phytoglyphic studies which largely enabled us to sort the hundreds of available specimens into groups which formed the basis of species descriptions. In what follows we now provide a resume of the sorts of information which can be obtained from phytoglyphic studies of the Corymbosae and which can be used as aids to species identification. All the photographic illustrations are light micrographs of stained leaf cuticles. Qualitative features of the leaf epidermis in Corymbosae A wide variety of phytoglyphic features 1s shown by different species of the Corymbo- sae. Vhe cuticular ornamentation of the subsidiary cells of the stomata may be very simple, as it is in the Eximiae (or yellow bloodwoods) and in, e.g. E. nesophila, or it may be very complex, as in e.g. E. latifolia and E. capricornia (Fig. 16). The ordinary, non- stomatal cells may contribute to the pattern of ornamentation (as they do in E. /atzfolia) or have little or no ornamentation. The illustration of E. datifolia was made from a speci- men of that species from Papua-New Guinea, and the pattern matches exactly that of specimens from Australia. Similarly the illustration of E. nesophila is from a specimen from Cape York Peninsula and the ornamentation 1s exactly the same as that of speci- mens from the Northern Territory and Western Australia. The pattern is a constant fea- ture of the species and within limits, does not vary over the entire range of the species or in cultivation. Moreover, it is constant over time, so that specimens collected during the last two centuries may be compared and identified with recently-collected specimens. As Mueller was aware, some eucalypts have no stomata on the upper surface of the leaf, or may have many fewer stomata on the upper surface than on the lower. Contrary to what Maiden wrote, these are constant characteristics for the species. For instance, E. polycarpa has scattered stomata on the upper surface of the leaf with a stomatal density of about | per square millimetre (Fig. 17), except near the margins and midrib where the density is about 3 per square millimetre. (Other, related species, have either no stomata on the upper surface or higher densities of stomata than E. polycarpa). The cap cells of oil glands have rather thin, unornamented cuticles. In the Corymbo- sae they are arranged in characteristic groups of 3 or more per oil gland and so are easily recognized in the light microscope. Some species, such as E. terminalis have large num- bers of groups of cap cells on both sides of the leaf; others (e.g. E. ollaris, E. opaca) have few or none (Fig. 18). There may be differences between upper and lower surfaces. For instance E. polycarpa has very few such groups on the upper surface, but large numbers on the lower surface. Similarly the actinocytic ‘giant stomata may be less common on the upper surface than on the lower; some species (e.g. E. opaca, E. centralis) have relatively few giant stomata on either surface of the leaf, while other species (e.g. E. terminalis) have many on both surfaces. We shall make further reference later on to ‘giant stomata’. Finally, the stomata of some species are in general larger than those of other species (Fig. 19). If studies of macroscopic morphology such as fruit shape and size, features of the flower, leaf shape and arrangement etc. suggest that two specimens are of the same species, we may expect that their phytoglyphic features will be identical. In Eucalyptus IT we have shown that the specimens which (wrongly) have been called E. perfoliata have features of the flowers which are identical with those of E. lamprocalyx. The fact that the phytoglyphic features of these specimens are identical with those of E. lamprocalyx (Fig. 20) supports the hypothesis of identity, and since the only validly-published name is that of E. lamprocalyx, it is under that name that we must now file specimens previously termed ‘E. perfoliata’. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 123 S25 7: E..terminalis Fig. 18. E. terminalis, with many groups of oil gland cap cells (asterisks) and E. opaca with only one. Arrow- head, giant stoma of E. terminalis. Scale bar: 500um. See also legend to Fig. 16. E.foelscheana “=. Y * / '\E.Rakad ee Eh Fig. 19. E. foelscheana and E. kakadu, to show differences in size of stomata. Scale bar: 50um. See also legend to Fig. 16. PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 124 THE ELASTIC-SIDED GUMLEAF My %, i E.lamprocalyx i Fig. 20. E. lamprocalyx (isotype) and E. ‘perfoliata: Identical phytoglyphs supporting identity of specimens. Scale bar: 50um. See also legend to Fig. 16. Pe gee ee ee SO 2 i lll chippendale - lenziana \ Fig. 21. E. chippendalei, with well-ornamented cuticle and E. lenziana with poorly ornate cuticle. Scale bar: 50pm. See also legend to Fig. 16. - w Views un i PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G. M. AND D. J. CARR 125 On the other hand, the cuticular ornamentation of closely related species, such as E. lenziana and E. chippendaler, may be quite different in appearance (Fig. 21). These differences may therefore be of very great diagnostic value, especially when the two species (the relationship between which may be evident, as they are in this case, from similarities of anumber of macroscopic features) occur together in the same region. These examples are intended to show how phytoglyphic features, taken together with comparisons of the macroscopic features of leaf, fruit and flower can be used to establish identity between specimens of the same species or, alternatively to establish that they are of different species. Since Mueller’s time, many publications have appeared testifying to the usefulness of such microscopic features of the leaves in taxonomy. Since palaeobotanists often have only fossilized leaves to work with, they have pursued the study of such features much more intensively than taxonomists of living plants. Unfortunately, all too few Australian botanists, other than one or two palaeo- botanists, have paid any attention to these features of either living plants or fossil leaves. For both Monocotyledons and Dicotyledons it is now, however, the general consensus that the pattern and shape of the leaf epidermal cells and features of their cuticles are species-specific and differ from one taxon to another (Barthlott and Ehler, 1977). Cell patterns involve the distribution of different types of epidermal cells and their positional relationships to each other and to the underlying tissues (e.g. of the veins) (Barlow and Carr, 1984). The presence and the distribution of idioblasts other than stomata (e.g. the cap cells of oil glands, emergent oil glands, hairs etc.) in the epidermis affects these patterns in species-specific ways. For instance, the first-formed stomata of the leaf are so-called ‘giant stomata. They arise near the margin and near the midrib, positions on the leaf primordia which remain fixed, relatively to the rest of the lamina, since marginal meristem activity in eucalypt leaf primordia ceases very early in develop- ment. Later on, some more giant stomata may be laid down in the intervening and expanding region of the lamina. In this region, the giant stomata may occupy positions near the centre of vein islets or over the junctions of smaller veins. These are also sites at which epidermal cells may differentiate to form the initials of oil glands. Oil gland initials may also arise early in leaf development near the margin and near the midrib. Thus in both situations, there is some interchangeability between the initials of oil glands and giant stomata, idioblasts which therefore compete, as it were, for preferred positions in the epidermis. On the other hand, neither oil glands nor giant stomata are a constant feature of the pattern and some species (e.g. E. opaca) exhibit a dearth of both in their leaf cuticles. We have already referred to ‘giant stomata’ as actinocytic, 1.e. they generally have a halo of numerous subsidiary cells so that they differ in appearance from the ordinary stomata. Such giant stomata were reported to be a feature of a number of Myrtaceae by Bandulska (1923; 1928-31), who found them on fossilized leaves of Myrtaceae in the Eocene of the London Clay deposits. Solereder (1908) who also reported the existence of giant stomata in some mangrove species referred to them as ‘water stomata. This is a misnomer. There is no evidence as far as we are aware, of a function of these stomata in secreting water. Nevertheless, some recent writers, including Stace (1965) have persisted in using the term ‘water stomata. Van Cotthem (1971) refers to them as hydathodes (in e.g. Buxus spp.). Van Wyck et al. (1982) also refer to ‘water stomata’ of the leaves of South African species of Eugenia. Napp-Zinn (1973-74) doubts the existence of giant stomata and declares his astonishment that Sitholey and Panda (1971) report on the giant stomata of Mangzfera indica and Limonia acidissima, without giving measurements of their actual size. Indeed, in terms of the length of the guard cells, the so-called giant stomata may, in some cases, not be larger than the ordinary stomata. Solereder reported ‘water stomata as either larger than or smaller than, the ordinary stomata. Nevertheless, PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 126 THE ELASTIC-SIDED GUMLEAF Mi) Wiad E. deserticola E. ferruginea “\ \ E.ferruginea (RRA iN Fig. 22. Giant stornata of E. maculata, E. deserticola and E. ferruginea, in comparison with ordinary stomata of these species. Scale bar: 50um. See also legend to Fig. 16. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 127 because of their larger complement of subsidiary cells, the giant stomata do stand out as different from — and their guard cells are indeed in most cases larger than — the ordinary stomata (Fig. 22). This is the case in many of the species of the Corymbosae. In addition, the cuticular ornamentation of the subsidiary cells of giant stomata is often less ornate or well-developed than that of the subsidiary cells of ordinary stomata. Quantitative aspects of phytoglyphic features of the Corymbosae We have already touched on two quantitative aspects of the phytoglyphic features of the leaves, vzz, differences in stomatal size between two species and the possibility of a very low stomatal density of the upper surface of the leaves in some species. Between related ‘heterogenous’ species (to use Mueller’s term) the density (number per unit area) of the stomata on the upper surface may be quite different. E. derbyensis, for instance has up to 90 stomata per square mm, as compared to the | per square mm in the related E. polycarpa. Frequency-spectra of the subsidiary cells The difference between giant and ordinary stomata in the number of subsidiary cells is not the only such difference. Even among the ordinary stomata some will have 3, some 4, some 5, etc. subsidiary cells. If one makes counts of the subsidiary cell comple- ment of a hundred stomata the percentage of 3s, 4s or 5s, etc. appears to be a repro- ducible characteristic of the species. For instance even a glance at a cuticle of a specimen of E. centralis shows that most of its stomata have 3 subsidiary cells (Fig. 23), whereas in a Fig. 23. E. centralis. All stomata in field each with 3 subsidiary cells, to compare with E. opaca, some stomata with 4, some with 5 subsidiary cells. Scale bar: 50m. See also legend to Fig. 16. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 128 THE ELASTIC-SIDED GUMLEAF similar cuticle preparation of the related E. opaca, there are more usually 4 or 5 sub- sidiary cells to each stoma. Before we consider the exciting possibilities of the diagnostic uses of such frequency-spectra of subsidiary cells we must first enquire into their constancy or otherwise in a single leaf, in different leaves of a single specimen, and between differ- ent specimens of a single species. Taking the latter first, we see (Table 1) that between 5 TABLE 1 Eucalyptus centralis 5 different specimens lower upper Subsidiary cells (%) 3 4 5 6 3 4 5 6 Specimen Ford 50 90 7 1 0 83 17 0 0 Carr 752 (type) 89 11 0 0 84.5 15 0.5 0 Jacobs 143 90 11 0 0 80 16 4 0 George 12954 84 15 1 0 84 15 1 0 Frith 49 86 12 2 0 79 18 3 0 Means 87.8 11.2 0.8 0 82.1 16.4 1, 0 x 0.066 2.929 = 0.312 0.427 = P>0.95 <0.98 n.s. (P<0.90) P>0.99 P=0.98 specimens of E. centralis there are no differences between percentage of 3s to a very high level of probability. The very much smaller percentages of 4s and 5s would require very large samples to be taken (perhaps 500 or 1000) to show whether or not the percentages are repeatable from specimen to specimen. The figures shown are based on counts num- bering between 100 and 200. Another feature is shown by the table: the upper surface generally has a lower percentage of 3s than the lower. The percentages of both 3s and 4s on the upper surface is the same in all the specimens, to a very high degree of prob- ability. The frequency-spectra of 6 specimens of E. centralis are shown in the graphs (Fig. 24). Evidently the spectra are highly reproducible from one specimen of a given species to another. The method of sampling to obtain the data was, using an oil-immersion objective and a stained, iverted cuticle preparation, to count the subsidiary cells of two or three stomata in a microscope field, then move to another field, repeating the process until over 100 counts had been obtained. By subsidiary cells, we mean all the cells which have a wall or part of a wall in common with one or both of the guard cells (see below, for a dis- cussion of this definition). If possible, the preparations for this purpose were taken from the middle third of the leaf. Will any leaf from a given specimen suffice for this pro- cedure? To test this, preparations were make from 5 leaves of a single specimen and counted. The data (Table 2) show that the 5 leaves yield frequency-spectra which are, to a very high degree of probability, identical. Again, the same sorts of differences between upper and lower surfaces are apparent, asin Table 1. There is a tendency for the spectra of the upper surface to be shifted along the abscissa, as compared with the lower. But this is not universally the case, as experience has shown us. If the middle portion of a leaf is not available, will it make a difference if we use some other part? To test this a single leaf was cut into 5 transverse strips, tip to base. Evidently (Table 3) there is no statistical difference, to a high degree of probability, between the tip, the middle and the base. But a word of caution is necessary. Both tip and base are narrower than the middle and in them the margin and midrib come closer PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G. M. AND D. J. CARR 129 TABLE 2 Eucalyptus orventalis Carr 763 (type) 5 different leaves lower upper Subsidiary cells (%) 3 4 5 6 3 4 5 6 Leaf 1 60.4 31.6 6.4 1.6 47.9 44.17 6.6 1.6 Leaf 2 57.89 32.46 7.29 led 49.19 42.74 8.1 0 Leaf 3 58.6 32.3 6.77 Da 48.62 AORS Ha) lO 0.9 Leaf 4 57.6 B72 6.4 0.8 48.8 47.4 Bills 0.78 Leaf 5 58.18 36.36 6.4 0 49.8 44.53 4.45 eZ. Means 58.53 33.178 7.74 — 48.78 43.84 6.47 - x? O.O805. OAPs Beer 0.0588 0.065 4.75559 P0599 BS 0295) E029 00295 IP S099 IP S0.99<0.98 n.s TABLE 3 Eucalyptus ollaris 5 zones, tip to base, of a single leaf of Carr 827 lower upper Subsidiary cells (% ) 3 4 5 6 3 4 5 6 Zone 1 (tip) 72 29u4 2.6 62.7 28.4 8.8 0 Zone 2 TORUS 24762 2S 0) 62.2 35.43 2.36 Zone 3 73 2e 3 0 64.6 27.4 1239620) Zone 4 70.37 26.85 Bild 0 63.41 31.45 Boe9 il, Zone 5 (base) 72.41 25.82 1.76 0 61.19 30.59 8.2 0 Means 72.17 299338 3.88 = 62.88 30.65 6.11 = x? 0.01825 0.19054 — 011129 11.9305 = P>0.9 P>0.99 P0989) > in. (IPOS) together. These latter are regions in which there are concentrations of giant stomata, which, with their unusually large number of subsidiary cells, could affect the com- parisons we wish to make. In addition, there might be ‘edge-effects’ on the frequencies of subsidiary cells even in ordinary stomata in the vicinity of the margin and midrib. These effects could be quite disturbing in especially narrow leaves (e.g. those of E. nelsoni and E. fordeana). To test for these edge effects we made use of leaves of E. opaca, which as we have already mentioned, have relatively few giant stomata. The stomata were sampled close to the margin and close to the midrib and the data compared with counts made half-way between those regions (Table 4). Evidently there is an appreciable edge-effect, especially on the upper surface, where the percentage of 3s is only two-thirds that of the stomata of the intervening region. The recommendation 1s, therefore, to avoid counting close to the margin and midrib. Use of the frequency-spectra in the Corymbosae We have made great use of this quantitative aspect of the phytoglyph. It is particu- larly useful in distinguishing between specimens of two species from the same general area. For instance, E. centralis (with a preponderance of 3s) is readily distinguishable (Fig. 24) from EF. opaca (Fig. 25), which occurs together with it in the same regions of central Australia and the Great Sandy Desert. The specimens used for these graphs PROG. LINN. SOC. N.S.W., 110 (2), (1987) 1988 130 THE ELASTIC-SIDED GUMLEAF 100 80 2 @ = 60 3 2 BH 3 o A 4 iS p ra o ZB 4 res Els © 5 , 49 Be a o7 a B6 Eliz 0 = Carr 752 lower Carr 752 upper Frith 49 lower Frith 49 upper Eucalyptus centralis percentage OSES NOW W percentage OSGN8 NOOR W Jacobs 143 lower Jacobs 143 upper percentage percentage OSES NOORW OBEN NOOhw Forde 50 lower Forde 50 upper George 12954 lower George 12954 upper Fig. 24 and Fig. 25 (opposite). Frequency spectra of E. centralis and E. opaca. Each separate graph represents a single specimen or the means of several specimens. On the left of each graph, the frequency-spectrum (f-s) of the lower surface, on the right the f-s of the upper surface. were drawn from widely separated regions, in the Northern ‘Territory and in Western Australia. Specimens of three other central Australian species, E. eremaea, and the mountain top mallees, E. nelsoni and E. fordeana, are also readily distinguishable by their frequency-spectra (Fig. 26). The Forrest specimen of E. fordeana was collected in 1883 by the explorer, John Forrest, on the summit of Mt Augusta in Western Australia, a locality very distant from the localities in central Australia from which the other specimen was obtained. Nevertheless, the frequency spectra match. The frequency spectra of widely disjunct specimens of three other central Australian species, E. symoni, E. australis and E. connerensis (Fig. 27) are also consistent and enable the specimens to be grouped unequivocally. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 131 80 percentage 60 o mn 3 g 3 4 5 4 ee o 40 El 5 A 6 a 6 Ee7 oO 7 20 : Ces opaca mean lower opaca mean upper Carr 808 lower Carr 808 upper Eucalyptus opaca @ is) @ oO mn = m : s m3 3 : 5 : = ff 5 ra) El 5 ‘= a M6 o M6 O 7 oO 7 ‘ Forde 66 lower Forde 66 upper Symon 9381 lower Symon 9381 upper percentage wo Oo ONES NON SW percentage OSES NOL W BS wwrers| ‘ : Forde 60 lower Forde 60 upper George 15668 upper TABLE 4 Eucalyptus opaca Carr 808 Edge effects: counts near the midrib, near the margin and in intermediate regions of the leaf lamina lower upper Subsidiary cells (%) 3 4 5 6 7 3 4 5 6 intermediate 68207 27-6 2.86 O95 O 62.39 34.19 3.4 0 near midrib 62.7 Bano 1.58 OO 39.27 48.2 9.82 2.68 near margin DIAM KOS a0) (),9) 0.9 38.68 94.72 6.6 0 Means 62.85 32.95 2.293 O.8) — 46.78 45.70 6.6 = x? 0.038 0.8687 0.006 - 5.2089 2.8989 = P>0.95 <0.98 n.s. P<0.90 P>0.99 n.s.(P<0.05) ns. (P<0.05) PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 UZ THE ELASTIC-SIDED GUMLEAF 60 50 GY Y =) 40 Y B > Y = Y 7 3 g Y 1 e ® 30 Y , § Y 2 Y : @ Y 3 %Y 6 5 Z 20 Yj o7 a Yj : yi K ‘ Gee ie G. : Maconochle 658 lower Maconochle upper Kalotas 452 lower Kalotas 452 upper Eucalyptus eremaea 100 80 > 60 Mg 3 3 = 4 c ; ° 5 3 5 40 6 o A 6 a oO 7 = (a) 7 mm 5 es Dunlop 2264a lower Dunlop 2264a upper Nelson 2249a lower Nelson 22498 upper Eucalyptus nelsonii 80 80 » © 60 S H 3 o = w 4 = 3 o < & ZB 4 e 40 fB 5 c 40 Gs oa : Ae gs g o7 20 ny o SS Vim y G 0 a ie, Forrest Mt Augusta lower Forrest upper 0 : S Ounlop Heauvitree lower Dunlop upper az Eucalyptus fordeana Fig. 26. Frequency spectra of E. eremaea, E. nelsoni and E. fordeana. In northwestern Western Australia, specimens of two species with somewhat over- lapping distributions, E. pyrophora and E. bynoeana are also readily separable by their frequency-spectra (Fig. 28). Again, in Queensland, specimens of two species, E. pocillum and E. capricornia, which occur together in a number of areas have quite different fre- quency spectra (Fig. 29). It is also to be noted that a Northern Territory specimen of E. capricornia (Lazarides 7094) has the same frequency spectra as the Queensland PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 133 80 80 a oO 60 bh o 40 percentage OSS NOOO SW percentage ONSS@ NOMA W 20 nN oO 0 7 Morecombe (WA) lower Morecombe upper Symon 2158 lower Symon 2158 upper Eucalyptus symonii percentage OSG NOW W percentage OSS NOOO SW Wilson 2480 lower Wilson 2480 upper 80 @ o 2 60 M3 D 6 3 S 4 = 4 S 5 ® BI 5 5 40 6 5 40 6 a Ez, a oO 7 20 20 0 Symon 9578 lower Symon 93578 upper Symon 2642 lower Symon 2642 upper Eucalyptus connerensis Fig. 27. Frequency spectra of E. symonit, E. australis and E. connerensis. specimens. This again confirms that, in general, the frequency spectra of widely dis- Junct specimens of a given species match each other. These quantitative aspects of the phytoglyph are thus of great help in grouping widely disjunct specimens of the same species as well as in discriminating between pairs of sympatric species. The frequency- spectra thus constitute an invaluable tool in dealing with species of the Corymbosae. Its potential value in dealing with other groups of eucalypts has still not been assessed, but we recently made some preliminary observations which showed that seedlings of E. paliformis and E. fraxinoides (Renantherae) have useful quantitative phytoglyphic differences. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 134 THE ELASTIC-SIDED GUMLEAF 50 ae a a 40 B 3 > 40 3 iS : = 4 < 30 5 ® 30 5 ct) =) o A6 L 6 S c-3) 220 O7 2 204 jal 7 Bs ] Hs 10 10 ZA) 0 Z 7am] Pyrophora mean/ lower mean/ upper KFK 7605 lower KFK 7605 upper Eucalyptus pyrophora & Y & £ Y counts 3 = o Y 4 ce 2 Y Hs 2 5 5 Y 6 a 7 j Y : Walcott lower Walcott upper Paton stop 7lower Paton stop 7? upper Y Bead : = Y ms s ; c Y 4 o 5 ° Y 5 2 6 3 Y 6 a Oo 7 a ZY o7 Be Y Barker lower Barker upper Fox & Brigden lower Fox & Brigden upper Fig. 28. Frequency spectra of (1) E. pyrophora, (2) E. bynoeana. Consideration of frequency-spectra in terms of development The basal number of subsidiary cells in the anisocytic stomata of the Corymbosae is three. We may consider that higher numbers arise from segmentation of the original 3 subsidiaries in the stomatal complex (Fig. 30). There are two possibilities for division of these initial cells. If each divides once, radially, the stoma will have 6 subsidiary cells; if only one, or only two, divide radially the number will be 4 or 5. Alternatively, one or more subsidiary cells may divide radially more than once, to give 7, 8 or 9 subsidiary cells. This is evidently more likely to occur during the development of giant stomata. The second possibility is that the proto-subsidiary cells divide, not radially, but peripherally. This would leave the subsidiary cell complement at its original number, if we consider the subsidiary cells as those which have a portion of cell-wall in common with one or both of the guard cells. In the Commelinaceae, for instance, the stoma may be surrounded by a series of rings of cells, forming a rosette which develops by successive peripheral divisions of 2 or 4 subsidiary cell initials (Tomlinson, 1969). We would use the term sub- sidiary cells for the innermost cells, those which actually share a wall with the guard cells. In connection with such cases, Napp-Zinn (1973) discusses the problem of PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G.M. AND D. J. CARR 135 percentage SON ann Ww percentage SOS8 nnht WwW Bynoeana mean mean upper lower Eucalyptus bynoeana 80 fo) Oo percentage > io) HOSS Nnns WwW percentage SEN nnhk Ww nN So Bynoe 2 lower Bynoe 2 upper percentage L o HES AnbawW SBS Daw percentage 20 Buncen ROWE aa BUN CEM UD Der Paton 42a lower Paton 42a upper definition of the term ‘subsidiary cell’, pointing out that many authors use the term not only for the innermost cells, as defined above, but also for what he terms ‘encircling cells’ (Kranzzelle) which may or may not owe their developmental origin to the single stomatal initial from which the rest of the stomatal complex is derived. Korn’s studies of stomata were directed, not to the numbers of subsidiary cells, but to the spacing and pattern of distribution of stomata in the epidermis. Discussing the helicocytic (sequentially spiral) divisions of the stomatal initials of Sedum stahli:i (Crassulacaeae), the last division of which produces the guard cells, surrounded by (in our definition) 3 or 4 subsidiary cells but, according to Korn, 6 subsidiary cells, Korn (1972) states that: ‘Regardless of which cells are true subsidiary cells, the methods employed in models discussed here used cell distances from a developmental aspect and not from the final position of cells. It can be suggested here that perhaps subsidiary cells of Sedum stahli are not subsidiary cells in the traditional sense but may serve as spacers to give ordered arrangements of stomata’. Whether or not the outer rings of cells in cyclocytic stomata, such as those of some genera of Combretaceae (Stace, 1963) and Myrtaceae (Bandulska, 1928-1951) have a PROG. LINN. SOG. N.S.W., 110 (2), (1987) 1988 136 THE ELASTIC-SIDED GUMLEAF 60 40 percentage top} Oo Shree SHS Nn bh WwW SOS nn Ss Ww pocillum mean lower mean upper Walker No 4 lower Walker No 4 upper Eucalyptus pocillum 100 80 SN anh WwW percentage SENB nn bk WwW Carr 1736 lower Carr 1736 upper SES DnLwW SOS AuhwW 40 percentage 20 — = 0 Bissett 684 lower Bissett upper Carr 149 lower Carr 149 upper Fig. 29. Frequency spectra of (1) E. pocillum, (2) E. capricornia. Lazarides 7094 is a specimen of the latter from Northern Territory; the others are from Queensland. physiological function in stomatal opening and closing like that of the innermost cells 1s, of course, unknown. Evidently the species-specific frequency-spectra of the Corymbosae must be genetically determined. This must also be the case for the rather rigid, almost mechani- cal sequences of divisions in many genera of Monocotyledons. It must be true also of families and genera of Dicotyledons in which the patterns of cell division of the stomatal complexes lead to specific and recognized ‘types’ of stomata, in which the numbers and arrangement of subsidiary cells, either 2, 3 or 4, are fixed and relatively invariant. Some of these patterns have been recognized as characteristic of whole families of flowering plants, e.g. the ‘cruciferous’, ‘ranunculaceous’ and ‘rubiaceous’ types. Because of their recurrence in many families, not only those after which they were named, these types are now Classified more objectively in terms of the number and arrangement of the sub- sidiary cells in the stomatal complexes. A considerable number of ‘stomatal types’ have been recognized (van Cotthem, 1970; Fryns-Claessens and van Cotthem, 1973; Dilcher, 1974; Wilkinson, 1979). Such classifications have led to the recognition of the widespread distribution of particular ‘stomatal types. For instance, the Magnoliales, PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G. M. AND D. J. CARR 137 Y Y, Ep Y Y m 2 m = Z Y 3 F ae @ 30 Y Y 5 2 El 5 o Y Y 6 5 6 a 20 Yy Y ol 7 a fla: 10 Ux Y Y Y mean upper Lazarides 7094lower Lazarides upper Eucalyptus capricornia 60 o GY = & 40 Y = LJ 2 Yee lie ° 4 ¢ a Y 4 © 95 ° Y 5 a M6 cy) Y 6 oO 7 20) Y El 7 10 0 n Simmons 4lower Simmons 4 upper 80 80 60 Y 60 2 Y 2 2 Y 3 2 B 3 4 Z 3 40 ZY a5 @ 40 5 5 Y 6 3 6 a Y 7 a (Oo) 7 20 Y 20 GE 0 a pos (0) oom Banks lower Banks upper Brass & White 121 lower Brass & White upper sensu ‘Takhtajan, have a nearly uniform occurrence of paracytic (1.e. ‘rubiaceous’) stomata, held by some writers to be a primitive type. In the Piperaceae, for instance, the stomata have been described as helicocytic and cyclocytic (van Cotthem, 1971). These relatively ‘standard’ patterns and numbers must be the products of sets of developmental steps, themselves the resultants of genetic programming by a relatively invariant seg- ment of the genome which is widely distributed in the species and genera of a number of families. This genetic programme determines the pattern and number of divisions giving rise to the stomatal complexes and therefore determines the number of subsidiary cells. Evidently the execution of the developmental steps involved may be subject to occasional aberration but, by and large, the genetically-determined end product will be predominant. In the Corymbosae, the genetic control of the frequency-spectra must also operate by determining the number and orientation of the cell divisions in the cell complexes of the stomatal initials, which are basically anisocytic. We may speculate that the cells, other than guard cells, of the complex have an initial capacity for cell division which 1s, within certain limits, determined genetically so that there is a statistical probability of a certain mean number of cell divisions, with less probability of fewer or more divisions. Moreover, the general orientation of these cell divisions, either predominantly radial, as in E. zygophylla and the related E. deserticola, or predominantly peripheral, as in PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 138 THE ELASTIC-SIDED GUMLEAF ns Fig. 30. Stomata to show radial subdivisions (1-3) and peripheral subdivision (4-6) of the initial subsidiary cells. All to the same scale. Scale bar: 50um. E. kakadu, must also be subject to genetic control. One interesting corollary of this view is that there must be genetic determinants which result in the patterns of distribution and of cell division in the stomatal complexes being, in general, different on the lower and the upper surfaces of the leaf. The differences between Monocotyledons, with their almost mechanical, deterministically-controlled patterns of cell divisions in the stomatal complexes and the Dicotyledons, many families and genera of which appear at first glance to have patterns of cell-division which are much more statzstically-determined, is paralleled by the developmentally-determined orientation of the guard cells in the Monocotyledons and the somewhat random arrangements in many (but not all) Dicotyledons. References BANDULSKA, H., 1923. — A preliminary paper on the cuticular structure of certain Dicotyledonous and Coniferous leaves from the Middle Eocene of Bournemouth. /. Linn. Soc. Lond. Bot. 46: 241-269. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 S.G. M. AND D. J. CARR 139 — , 1928/31. — On the cuticles of some recent and fossil Myrtaceae. J. Linn. Soc. Lond. Bot. 48: 657-671. BaRLow, P. W., and Carr, D. J., (eds), 1984. — Positional Controls in Plant Development. Cambridge: Cambridge University Press. BARTHLOTT, W., and EHLER, N., 1977. — Raster Elektronenmikroskopie der Epidermis-Oberflachen von Spermatophyten. Tropische und Subtropische Pflanzenwelt. 19, Mainz.Akad.Wiss.u.d.Literatur. Wiesbaden: Steiner. BLAKE, S. T., 1953. — Botanical contributions of the Northern Australia Survey. I. Studies on Northern Aus- tralian species of Eucalyptus. Aust. J. Bot. 1: 185-352. CarR, D. J., and CARR, S. G. M., 1978. — Origin and development of stomatal microanatomy in two species of Eucalyptus. Protoplasma 96: 127-148. , and , 1980a. — Eucalyptus stomata with occluded anterior chambers. Protoplasma 104: 239-251. , and ——, 1980c. — The Lehmannianae: a natural group of Western Australian eucalypts. Aust. J. Bot. 31: 629-643. , and ——, 1985. — Eucalyptus I. New or little-known species of the Corymbosae. Canberra: Phytoglyph Press. , and , 1987. — Eucalyptus II. The rubber cuticle and other studies of the Corymbosae. Canberra: Phyto- glyph Press. CaRR, S. G. M., 1972. — Problems of the geography of the tropical eucalypts. In D. WALKER, (ed.), Bridge and Barrwer, the natural and cultural history of Torres Strait: 153-181. Canberra: A.N.U., Dept Biogeogr. and Geomorph. Publ. BG/3. , and CarR, D. J., 1976. — Identification of a eucalypt fragment, based on anatomy of leaf and stem. Proc. Roy. Soc. Vict. 88: 77-82. , and , 1979. — An unusual feature of stomatal anatomy in certain taxonomically-related Eucalyptus spp. Ann. Bot. 44: 230-243. ——,, MILkovits, L., and Carr, D. J., 1971. — Eucalypt phytoglyphs: the microanatomical feature of the epidermis in relation to taxonomy. Aust. J. Bot. 19: 173-190. CUTLER, D. F., ALVIN, K. L., and PRICE, C E., (eds), 1982. — The Plant Cuticle. Linn. Soc. London, Symp. No. 10. London: Academic Press. DILCHER, D., 1974. — Approaches to the identification of angiosperm leaf remains. Bot. Rev. 40: 1-157. FRYNS-CLAESSENS, E., and VAN COTTHEM, W., 1973. — A new classification of the ontogenetic types of stomata. Bot. Rev. 39: 71-138. Korn, R. W., 1972. — Arrangement of stomata on the leaves of Pelargonium zonale and Sedum stahli. Ann. Bot. 36: 325-333. MAIDEN, J. H., 1909-1928. — A critical Revision of the Genus Eucalyptus. Vols I-VIII. Sydney: Government Printer. MarTIN, H. A., and JUNIPER, B. E., 1970. — The Cutzcles of Plants. London: Edward Arnold Ltd. METCALFE, C. R., 1983. — Secretory structures: Cells, cavities and canals. Jn C. R. METCALFE and L. CHALK, (eds), Anatomy of the Dicotyledons. 2nd edition, II: 64-69. Oxford: Clarendon Press. MUELLER, F. VON, 1874. — Observations on new vegetable Fossils of the auriferous Drifts. 2nd decade. Geological Survey of Victoria. Melbourne: Government Printer. ——,, 1879-1884. — Eucalyptographia, a descriptive Atlas of the Eucalypts of Australia etc. Melbourne: Government Printer. Napp-ZINN, K., 1973. — Anatomie des Blattes. II. Angiospermen, A.I. Encyclopaedia of Plant Anatomy, viii (2A). Berlin, Stuttgart: Gebrider Borntraeger. NEVILLE, A. C., 1963. — Daily growth layers in locust rubber-like cuticle influenced by an external rhythm. J. Ins. Physiol. 9: 177-186. SITHOLEY, R. V., and PANDEY, Y. N., 1971. — Giant stomata. Ann. Bot. 35: 641-642. SMITH, H. G., 1908. — On the elastic substance occurring in the shoots and young leaves of Eucalyptus corymbosa and some species of Angophora. J. Proc. Roy. Soc. N.S.W. 42: 133-146. SOLEREDER, H., 1908. — Systematic Anatomy of the Dicotyledons. Transl. L. A. BOODLE and F. E. FRITSCH: revised D. H. ScorT. Oxford: Clarendon Press. STAcE, C. A., 1965. — Cuticular patterns as an aid to plant taxonomy. Bull. Brit. Mus. (N.H.) Bot. 4: 1-78. TOMLINSON, P. B., 1969. — Anatomy of the Monocotyledons. 111. Commelinales-Zingiberales. Oxford: Clarendon Press. VAN COTTHEM, W. R. J., 1970. — A classification of stomatal types. Bot. J. Linn. Soc. Lond. 63: 235-246. ——, 1971. — Vergleichende morphologische Studien tber Stomata und eine neue Klassifikation ihrer Typen. Ber. dtsch bot. Ges. 84: 141-168. VAN WYCK, A. E., ROBBERTSE, P. J., and KOK, P. D. F., 1982. — The genus Eugenia (Myrtaceae) in southern Africa: the structure and taxonomic value of stomata. Bot. J. Linn. Soc. Lond. 84: 411-56. WELCH, M. B., 1923. — The secretory epidermal cells of certain eucalypts and angophoras. j. Proc. Roy. Soc. N.S.W. 57; 218-228. WILKINSON, H. P., 1979. — The plant surface (mainly leaf). Part I Stomata. In C. R. METCALFE and L. CHALK, (eds), Anatomy of the Dicotyledons. 2nd edition, I: 97-117. Oxford: Clarendon Press. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 140 THE ELASTIC-SIDED GUMLEAF ADDENDUM After delivery of the lecture, the president of the Society, Dr Peter Martin, kindly supplied us with copies of pages of two books on organic chemistry by Professor John Read F.R.S. in which reference is made to Henry G. Smith’s discovery of the rubber cuticles of certain bloodwoods and species of Angophora. The works are: A Textbook of Organic Chemistry. Historical, Structural and Economic, 3rd ed., London: G. Bell & Sons Ltd, 1948, p. 615, and A Direct Entry to Organic Chemistry, London: Methuen & Co. Ltd (Home Study Books), 1948 reprinted 1953, p. 227. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western Blue Mountains P. GLASBY, P. M. SELKIRK, D. ADAMSON, A. J. DOWNING and D. R. SELKIRK Gtiassy, P., SELKIRK, P. M., ADAMSON, D., DOWNING, A. J., & SELKIRK, D. R. Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western Blue Mountains. Proc. Linn. Soc. N.S.W. 110 (2), (1987) 1988: 141-158. Fire is an important factor in controlling the distribution of Eucalyptus oreades within its natural range in the western Blue Mountains. The species which is fire sen- sitive relies on seeding rather than resprouting after fire. In the western Blue Moun- tains, E. oreades produces annual rings in the wood, allowing the age of a tree and dates of fire damage to it to be determined. Stands of E. oreades thus allow establishment of a fire chronology in localized areas. In uniform-aged young stands which are undergoing self-thinning the suppressed individuals set seed before the dominant individuals. Trees which survive to become mature are less susceptible to fire damage, developing a bark skirt at the base which protects them from ground-fire damage. The trees enter the high fire risk period of summer with substantial seed reserves stored in capsules in the canopy. Wind-throw is a major cause of death of old trees, already fire- and/or termite-damaged. P. Glasby, P.M. Selkirk, D. Adamson, A. J. Downing and D. R. Selkirk, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109; manuscript recerved 12 May 1987, accepted for publication 19 August 1987. INTRODUCTION Eucalyptus oreades R. T. Baker, (Blue Mountains Ash), also known as smooth-barked Mountain Ash or White Ash, occurs on plateaus, ridges and in gullies in the Blue Mountains of N.S.W. It is valued locally not only as a beautiful tree but as a species of economic importance. Pole-sized saplings are logged in Newnes State Forest to supply pit props for coal mining. The Explorers’ Tree at Katoomba, marked by Blaxland’s party when making the first crossing of the Blue Mountains in 1813 is E. oreades (Baker, 1919). The vegetation of the Blue Mountains has been little studied despite the spectacu- lar topography of the region and its proximity to the vast urban area of Sydney. For example, E. oreades is not even listed in a recent review of wet sclerophyll eucalypts on the east coast of Australia (Ashton, 1981). Apart from the original description of E. oreades (Baker, 1889) and more recent descriptions, with maps, of its Australian distribution (Boland et al., 1984), no botanical publications deal in any detail with the species. Vegetation descriptions of the Mt Wilson area (Brough et al., 1924; Petrie, 1925) include associations which contain E. oreades and recognize that aspect and altitude are im- portant factors for its occurrence (Pidgeon, 1938). A vegetation classification of the western region of Sydney (Forster et al., 1977), and the unpublished Forestry Com- mission 1:25,000 map of Newnes State Forest No. 748, set out its general distribution in the Blue Mountains. Following studies on the ecology of wet sclerophyll species, particularly E. regnans, in Victoria and Tasmania (Ashton, 1958; Gilbert, 1959; Cunningham, 1960; Jackson, 1968), a picture emerged of the interdependence of tall eucalypt forest and fire, a picture which was not generally recognized by early workers on the forests of eastern New South Wales (Brough et al., 1924; Petrie, 1925; Pidgeon, 1937, 1938, 1940; Beadle, 1954, 1962). The response of EF. oreades to fire is well known in general terms to local staff of the PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 142 BLUE MOUNTAINS ASH Forestry Commission and the National Parks and Wildlife Service but no information has been published on its response to fire or on its general biology. As the urbanized area in the Blue Mountains expands rapidly, particularly along narrow ridges with an intrinsic high fire risk to property, there is increasing demand to minimize that risk by frequent burning, with little knowledge of its effects on the vegeta- tion in general or on certain species in particular. Present-day attempts to manage fire in whole National Parks and in urban bushland areas represent a major change from earlier, more laissez-faire, approaches to fire control, an approach to fire which E. oreades survived. With the present trend towards planned intervention in previously un- managed fire regimes, detailed information on the biology of the plants involved is clearly needed by land managers. An additional threat to E. oreades is the discovery that the Newnes Plateau and adjacent high altitude areas contain huge deposits of sand and clay suitable for deep open-cut mining (Pecover, 1984; Anon., 1984). Implications for the landscape and vegetation of the western Blue Mountains region are severe. This paper describes aspects of the distribution and life history of E. oreades which help to explain its occurrence in the fire-prone landscapes of the western Blue Mountains. DISTRIBUTION General Eucalyptus oreades occurs in a number of disjunct populations within the latitudes 28°15’S to 34°30’S (Fig. 1). The largest population is in the Blue Mountains. Other stands occur on the escarpment inland from Port Macquarie, in the Gibraltar Range between Grafton and Glen Innes, in the Binna-Burra—Springbrook— Mt Warning area near the New South Wales— Queensland border, and near Tenterfield (Boland et al., 1984). The altitudinal range lies between about 700 and 1200 metres. The disjunct distribution could indicate a former, more continuous, distribution. Blue Mountains In the Blue Mountains, E. oreades occurs on sands and clayey sands derived from Triassic sandstone parent rock in the Katoomba and Grose soil associations (Forster e¢ al., 1977). It also occurs on alluvial and colluvial sands in valleys and deep river gorges, on cliffs, and on sandstone soils whose texture is influenced by deep, red clay loams der- ived from the basalt caps at Mt Wilson. It would appear that soil depth or type does not explain its distribution in the Blue Mountains. A striking feature of the species’ distribution in the Blue Mountains 1s its relation- ship to the spectacular topography. It occurs rarely on exposed ridges but is common on steep sheltered slopes with a southerly aspect and around the heads of valleys facing south and east, on cliffs and cliff ledges, and along creek beds (Fig. 2). In these locations it can occur as individuals, particularly on cliffs, or as dense stands in sheltered valleys (in wet sclerophyll or tall open-forest). On the gentler slopes of the plateaus and ridges extending west from Katoomba to the Newnes Plateau, mature trees develop spreading crowns with short boles and branches low down on the trunk. Such trees have been kept as specimen trees in the towns of the upper Blue Mountains. They also occur as widely spaced individuals in mixed eucalypt open-forest associations (dry sclerophyll). By con- trast, trees growing in crowded stands on steep sheltered slopes develop straight tall trunks, topped by a simple domed canopy. These form stands of tall open-forest (wet sclerophyll). Trees growing on ledges and in joints on cliffs develop tall straight, often non-vertical, trunks. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 P. GLASBY ETAL. 143 (b) 150° 15'E Newnes State Forest Lithgow 33°30'S National Park £\ Mt.Wilson Grose (Valley Mt Bell Mt. | e Victoria Blackheath CD Katoomba Fig. 1. a. General distribution of E. oreades (after Boland ef al., 1984). b. Location of sites in western Blue Mountains mentioned in text. More detailed locations specified in some diagrams. Measured Transects Fig. 3 is a downslope section through a tall open-forest of E. oreades at Evans Look- out near Blackheath. This site (AB) is typical of dense stands which occur on south-to- east-facing slopes of steep valleys (Fig. 2b), particularly near the main drainage lines. The site is below the rim of the plateau and above the first main line of cliffs within a tributary valley of Greaves Creek. An open woodland of eucalypts including E. pzperita (Peppermint) and E. siebert (Black Ash) grows on the plateau. Closed forest (rainforest) of mainly Ceratopetalum apetalum (Coachwood) and Doryphora sassafras (Sassafras) occupies the lower parts of the valley below the first main line of cliffs. At this site Eucalyptus oreades occupies the typical position of wet sclerophyll tall open-forest: relatively sheltered sites located between the even more sheltered rainforest vegetation below and the more exposed woodland vegetation on ridges and plateaus. Beneath the E. oreades canopy 1s an open sparse intermediate canopy of Acacia elata (Cedar Wattle) and a lower dense canopy dominated by Callicoma serratifolia (Black PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 144 BLUE MOUNTAINS ASH Fig. 2. a. Two large spreading Eucalyptus oreades trees on an exposed ridge, Gordon Lookout, Leura. b. Dense stand of E. oreades at the head of a south-facing gully near Evans Lookout, Blackheath (adjacent to transect at site AB, Fig. 3). c. Eucalyptus oreades trees on cliff face near Govetts Leap, Blackheath. E. oreades dere { dead E. cypellocarpa a E. piperita WS E. sieberi t C. apetalum Ee) A.elata upper surface LN of shrub layer @ 1 Fig. 3. Transect at site AB (1981) at Evans Lookout, Blackheath, showing distribution of E. oreades at the head of a south-facing valley, between dry sclerophyll woodland on plateau surface and rainforest in lower cliff- bounded valley. PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 P. GLASBY ETAL. 145 Wattle). Dense clusters of ferns such as Sticherus lobatus and litter accumulations ensure very low light levels at soil level. Fig. 4 presents similar data for two other valley slopes which face between east and south. Site CD is on an east-facing slope of the linear N-S valley of Govetts Leap Brook in an area dominated by woodland, shrubland and heath. Occasional tall trees of Blue Mountains Ash occur near the creek. Site EF, in the upper Bowens Creek Valley, is simi- lar in situation to AB but EF was burnt in a severe fire in December 1979. Severe fire has probably not affected AB since 1959. In each site E. oreades clusters on sheltered moist slopes at the head of steep south-facing creeks beneath the scarp of a distinct ridge or plateau. / _oreades ‘\ live | dead . piperita NS . sieberi t . radiata . mannifera TI . apetalum upper surface of shrub layer Fig. 4. Transects at sites CD and EF (1981). CD near Govetts Leap, Blackheath, shows live and fire-killed E. oreades in a small isolated stand last affected by fire in 1959. The site was again burnt in 1982 (see text). EF isa site where almost all mature trees in a large stand were killed by severe fire in December 1979. Extensive seedling establishment has since occurred. Site GH (Fig. 5) traverses an asymmetric valley in the headwaters of Greaves Creek between Medlow Bath and Blackheath, where the Blue Mountains plateau is only slightly incised. Again, trees of E. oreades cluster on the east-facing steeper slope but widely spaced mature individuals occur in the woodland on the gentle slopes of the plateau. These tall trees project well above the other eucalypts and are usually associated with several small living or dead offspring. Although the sites in Figs 3 and 4 contain trees of widely different size and age, similar individuals are often clustered. The patchy nature of the stands suggests episodic death and regeneration in response to fire as the causal process. PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 146 BLUE MOUNTAINS ASH EFFECT OF FIREON STAND STRUCTURE Baker (1919) noted and we have confirmed that the wood of E. oreades grown in the Blue Mountains contained annual rings. Blue Mountains Ash also records fire damage in the wood where the vascular cambium is killed. Using fire scars dated from tree rings, a fire chronology was obtained for the sites AB, CD and EF and the ages of trees of various sizes were determined. Fig. 6a-c shows frequency distributions for tree size and the age and fire history of individual trees from these sites based on tree rings. The effect of fire is shown clearly at EF where the severe 1979 fire killed all trees below 0.5m stem diameter and many of those larger. These trees had survived fires in 1975, 1966 and 1959. At EF the survivors after the fire were a few large trees located downhill near the boundary of the closed forest. A swarm of seedlings established in the area occupied by the fire-killed trees and by December 1985 were up to 4m tall and sufficient in numbers to replace the earlier stand despite below average rainfall in 1981-1982 and several months of drought after the fire. east 0 200 400 600 800 1000m — valley profile —-—~— general level of canopy E. oreades Fig. 5. Transect at site GH (1980), across upper Greaves Creek valley, between Medlow Bath and Blackheath, on the west of and parallel to electricity transmission line. At Govetts Leap Brook (site CD) the swarm of small trees was killed in a severe fire of late 1982. The one large, mature tree survived but the fire killed half of its canopy and half of its vascular cambium to a height of 1.5m, severely weakening it. Since this fire, only 4 seedlings (up to 0.5m tall in December 1985) have become established. At the Evans Lookout site (AB) a fire in 1959 was followed by the establishment of a cohort of trees dating from about 1960. This fire burnt patchily in the valley leading to localized death of trees, scarring of others, and the regeneration of new plants. A later less severe fire occurred in 1969. Each site shows that fire easily kills young trees of E. oreades and severe fire, particu- larly through the canopy, easily kills mature trees. Following each fire there is usually recruitment of a new cohort of plants from seed, leading to the development of a mosaic of even-aged stands of different ages, each stand in its own burn patch. Observation of fire-damaged trees shows that fire kills the live bark and vascular cambium of trunk and PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 P. GLASBY ETAL. 147 branches and that the surviving parts of damaged trees have little capacity to resprout. This sensitivity of individual trees of E. oreades is in marked contrast to the vigorous resprouting after severe fire of most other eucalypts in the region. 40 F K © F F ---- F —---- KK a (a) 20 —_ Tae TT a0 Amar Ra psa ala nC eT 1980 1920 0 0 50 60 ———-® 2 F-Fire scar roa " K-Kino vein (indirect evidence uj 40 1980 | of possible fire) (b) = -Presumed date of = |, establishment from seed Zz 00 i—-Date of establishment uncertain; tree centre decayed 0 0 50 F F K KKKKK___» ue sera easy : BG ! i maa, FAST ARTS TED LSS ETC | fe SEEDLINGS a (c 1 |! post79-80fire 1980 1960 1920 10 pre79-80fire 0 50 100 STEM DIAMETER (cm) Fig. 6. Frequency distribution in mid-1980 of trees of various diameters with fire history of particular trees inferred from examination of annual rings and fire scars. a. Site AB Evans Lookout. b. Site CD Govetts Leap Brook. c. Site EF Bowens Creek valley. Establishment of large numbers of plants following a fire leads to strong intra- specific competition and the development of a wide range of size within an even-aged stand. The dominant individuals grow rapidly; the suppressed individuals grow slowly and most die. Fig. 7 shows size classes in such a stand about 12 years after a fire. ‘Trees ranged in height from about 2m to 8m. The most suppressed trees had very little (about 2cm) new growth per shoot, bore capsules from two years, buds and very few leaves. The dominant trees in the stand had extensive new shoot growth (about 30cm per shoot), bore no capsules or buds, and had a crown of dense foliage. After a further four years self-thinning had removed the smallest trees. PROG. LINN. SOC. N.S.W., 110 (2), (1987) 1988 BLUE MOUNTAINS ASH 148 6 000 96 000 6& 000 F 000 000 $6 OOO TTT (og <)¢ 201 6 000 8c 000 &P 0009 000 I 000 96 000 09 (0§ <)Z24L & 000 €9 000 1¢ 000 G 000 6 000 OT 000 %6 (001 <) 1 994 poos sopnsdeo sofnsdeo sofnsdeo sojnsdeo aqnsdva a[qeia IV Ad ajding usa15 spng (stead ul ase) Jad paas Ayn SNIGIEMAN 904) Jod feo J, $a] SOPBIIO “TY papjaf-purm ainjout ary) fo sardouns ay) U2 pasojs paas pun ‘saynsqvo ‘spnq fo saqunu fo saquensqns pajunos uLoLf sayvUcrjsy | TEV L, 110 (2), (1987) 1988 N.S.W., LINN. SOC PROC P. GLASBY ETAL. 149 SEED PRODUCTION IN SUPPRESSED TREES A striking feature is that the suppressed trees in the 12-year-old stand produced capsules whereas the dominant trees did not (Fig. 7). Each capsule contained about 3 viable seeds, a similar number to that of mature trees (Table 1). The total number of viable seeds per suppressed plant (average of about 350) was low because of the small number of capsules on each tree although the total number of seeds per unit ground area in the stand was approximately 36,000 per 100m?, a substantial fraction (about 4 to %) of that found in a stand of mature trees because of the large number of suppressed plants. The early and effective reproductive behaviour of suppressed trees in a crowded uniform-aged stand constitutes a hedge against a severe fire in the period before the dominant individuals reach maturity. Diversion of resources from vegetative to reproductive growth in the suppressed trees guarantees their early death but provides an early seed bank at that particular site. This behaviour 1s an effective insurance against a severe fire following between about 10 and 25 years after an earlier one. SOF Trees a with capsules 60 C] without capsules —~ less 2-5cem greater than 2cm than 5cm DIAMETER NUMBER PER 100m? Fig. 7. Size distribution of a dense young stand of E. oreades regenerated after clearing and fire in 1969. Draw- ings are representative of three size classes. Stem diameter measured 10cm above ground level. Stand located at the head of a south-facing valley on the southern edge of the Katoomba Airfield (Medlow Bath). SEED AND LITTER PRODUCTION IN MATURE TREES Mature E. oreades trees bore three distinct age classes of capsules, as well as develop- ing buds (Table 1). The outermost growing tips of branches bear new vegetative growth and young flower buds which open in summer. Behind these, within the mature leafy PROC. LINN. SOC. N.SW., i10 (2), (1987) 1988 150 BLUE MOUNTAINS ASH area of the branch, are the green capsules from the previous summer’s flowering. Further back is an area of small branches usually devoid of leaves which bear purple capsules which are two years old. Still further back on the branches are grey capsules borne either on persistent dead twigs or on the main branch itself. These grey capsules may be three years old or older. Viability of seeds from the three colours of capsules is shown in Table 2. Only 2% of seeds from green capsules collected in August 1980 were viable, but by October, all ages of capsules contained viable seed. In all capsules larger, heavier seeds were more likely to be viable than smaller, lighter ones (Table 2). E. oreades therefore enters the high risk summer fire period with large amounts of viable seed stored in the canopy (Table 1). TABLE 2 Percentage germination, and weight per seed, of three length classes of seeds from three capsule ages harvested in October 1980 from storm-felled trees Germination was tested on four replicates of 25 seeds. The standard errors are given in brackets Length of Capsule seed (/) Toisermination PISTERE seed colour longest axis weight (mg) (mm) green 1.8 SE ee cue usyueg eiISOUIOH rt % 001 0) aaoibueyy aa — api) Moy Sat peop l epit YBIH or aN): (ES “suouttoads aptuaant SayBOIPUI YsaIsy “YeIq ul uMoyYs st suauttoads peap jo uonsodoid ‘ureiSetp jo yoy doi uy ‘sofdures Jo uoneo0] SOAIS LIOT] PoleIIOsse Sutmoys ayyoid a10ys -IR2N “%OFS'(99S = L) ¢ a[dures ul pue °% cy st(6G6zZ = L) Zz a[dures Ul °%BE SI(L8S = L) , e[dures ut suouttoads peap jo uonsodoig “anptsail poysem yo 099¢ sad spenptarput yo jaquinu [e101 st], “(T “Sly yo y Artjeso]) Sure py WIOg “WHY ISA YINOS UL QUOZ [BPN UI SapoOde.}sSO jo uonnqimsiq ¢ 327 N.S.W,, 110 (2), (1987) 1988 LINN, SOC. PROC, 164 OSTRACODES FROM PORT HACKING 0 50 100% 0 Sample 1 T=193 0) 50 100% 2 Sample 2 T=610 4m Osticythere baragwanathi (CHAPMAN & CRESPIN, 1928) Paracytheroma sudaustralis (McKENZIE, 1978) Callistocythere dorsotuberculata paucicostata YASSINI & JONES Actinocythereis (Ponticocythereis) militaris (BRADY, 1866) Cytherura portuswelshpoolensis HARTMANN, 1980 Loxoconcha australis BRADY, 1880 Xestoleberis cedunaensis HARTMANN, 1980 Leptocythere hartmanni McKENZIE, 1967 Pectocythere portjacksonensis McKENZIE, 1967 Hemicytheridea hiltoni Nn. sp. Microcytherura aestuaricola HARTMANN, 1980 Callistocythere puri McKENZIE, 1967 Callistocythere sp. nov. Semicytherura illerti sp. nov. “Hiltermannicythere’ bassiounii HARTMANN, 1981 Parakrithella australis MCKENZIE, 1967 Echinocythereis melobesioides BRADY, 1880 Australimoosella lauta (BRADY, 1880) Paradoxostoma augustensis HARTMANN, 1979 Xestoleberis chilensis austrocontinentalis HARTMANN, 1978 Caudites litusorienticola HARTMANN, 1981 Actinocythereis (Ponticocythereis) ichthyoderma (BRADY,1890) Loxoconchella pulchra McKENZIE, 1967 Loxoconcha sp. nov. Pectocythere sp. (Ceduna 120) HARTMANN, 1980 Aglaiella dietmarkeyseri HARTMANN, 1979 Bairdoppilata sp. Procythereis (Serratocythere) kerguelenensis (BRADY, 1880) High tide Low tide Posidonia australis Gracilaria sp. — Ectocarpus silicolosus Sargassum sp. 2 1 po AO) individuals Procythereis (Serratocythere) densuireticulata HARTMANN, 1981 Quadracythere obtusalata BRADY, 1880 Fig. 4. Distribution of ostracodes in profile off jetty at Cronulla Sailing Club, Gunnamatta Bay (locality B of Fig. 1), in marine zone of circulation. T is total number of individuals per 50cc of washed residue. Proportion of dead specimens in sample 1 is 35% (T = 193) andin sample 2 is 67% (T = 610). Nearshore profile showing associated floras gives location of samples. In top left of diagram, proportion of dead specimens is shown in black. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 AJW/DMM/GD962 I. YASSINIANDA. J. WRIGHT 165 neither illustrated nor mentioned in the text. Australimoosella lauta (Brady), Pectocythere sp. (Ceduna 120), Procytherers (Serratocythere) densuireticulata Hartmann, P. (S.) kerguelenensis (Brady), Quadracythere obtusalata (Brady) and Loxoconchella pulchra McKenzie are not illustrated. All specimens are deposited at the Australian Museum; AMP refers to specimens in the Recent Crustacea catalogue. All photographs are by SEM,and each is provided with a linear scale. SYSTEMATICS Suborder PLATYCOPIDA Family BAIRDIIDAE Brady, 1883 Genus Bairdoppilata Corywell, Sample and Jennings, 1935 Bairdoppilata sp. (AM P36478) Only two isolated valves of this form were encountered in Gunnamatta Bay and are not illustrated. Family CYTHERIDAE Baird, 1850 Genus Microcytherura Muller, 1894 Mucrocytherura aestuaricola Hartmann (Fig. 5G-T) (AM P36493) 1980 Microcytherura aestuaricola Hartmann, p.117, pl.3, figs 7-13 Family OSTICYTHERIDAE Hartmann, 1980 Genus Osticythere Hartmann, 1980 Osticythere baragwanatht (Chapman and Crespin) (Fig. 5M,N) (AM P36494) 1928 Cythere baragwanathi Chapman and Crespin, p.126, pl.10, fig. 65a,b 1980 Osticythere reticulata Hartmann, p.119, pl.4, fig. 7-18 1984 Osticythere reticulata Hartmann; McKenzie and Pickett, p.236, fig. 4, R-U 1986 Osticythere baragwanathi (Chapman and Crespin); McKenzie, p.107 Family LEPTOCYTHERIDAE Hanai, 1957 Genus Leptocythere Sars, 1928 Leptocythere hartmanni (McKenzie) (Fig. 7K,L) (AM P36489) 1967 Callistocythere hartmanni McKenzie, p.81, pl.12, fig. 5 1978 Leptocythere hartmanni (McKenzie); Hartmann, p.79, figs 101-107 1980 Leptocythere hartmanni (McKenzie); Hartmann, p.123, pl.5, figs 15,16,18,19 1984 Callistocythere hartmanni McKenzie; McKenzie and Pickett, p.239, fig. 5Y Genus Callistocythere Ruggieri, 1953 Callistocythere dorsotuberculata paucicostata Yassini and Jones (Fig. 7M,N) (AM P36479) 1987 Callistocythere dorsotuberculata paucicostata Yassini and Jones, p.27, pl.2, figs 3-4 PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 166 OSTRACODES FROM PORT HACKING Fig. 5. A-F, Hemicytheridea hiltoni Yassini sp.nov. A, LV with details of posterior hinge teeth and sockets; B, LV internal view; C, RV external view, holotype; D, Ventral view of carapace; E, RV external view; F, Details of ornamentation. Sample 2, Port Hacking, Gunnamatta Bay. G-I, Microcytherura aestuaricola Hartmann, 1980. G, LV external view; H, Details of ornamentation; I, LV internal view. Sample 2, Port Hacking, South West Arm. J-L, ‘Hiltermannicythere’ bassiounit Hartmann, 1981. J, RV external view; K (same scale as J), LV external view; L, Details of ornamentation. Sample 2, Port Hacking, South West Arm. M, N, Osticythere baragwanathi (Chapman and Crespin, 1928). M, LV external view; N, RV internal view. Sample 3, Port Hacking, South West Arm. O, P, Actinocythereis (Ponticocythereis) militaris (Brady, 1866). O, LV external view; P, internal view. Sample 3, Port Hacking, South West Arm. Q-S, Loxoconcha variolata Brady, 1878. Q, LV external view; R, LV dorsal view; S, RV internal view. Sample 1, Port Hacking, South West Arm. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 1967 1980 1978 1980 1967 1980 1984 13866 1880 1967 1976 1984 1890 1986 1880 1976 1978 Ia) 1980 I. YASSINI AND A. J. WRIGHT 167 Callistocythere purr McKenzie (Fig. 70) (AM P36480) Callistocythere purr McKenzie, p.81, pl.12, fig. 2; text-fig. 32 Callistocythere purr McKenzie; Hartmann, p.124, pl.7, figs 3,6 Family CYTHEROMIDAE Elofson, 1939 Genus Paracytheroma Juday, 1907 Paracytheroma sudaustralis (McKenzie) (Fig. 7C,D) (AM P36495) Cytheroma sudaustralis McKenzie, p.178, figs 30,35-42 Paracytheroma sudaustralis (McKenzie); Hartmann, p.128, figs 51-56 Genus Parakrithella Hanai, 1959 Parakrithella australis McKenzie (Fig. 7G) (AM P36497) Parakrithella australis McKenzie, p.72, pl.26 fig. N-O Parakrithella cf. australis McKenzie; Hartmann, p.129 Parakrithella australis McKenzie; McKenzie and Pickett, p.236, fig. 4G-H Family TRACHYLEBERIDIDAE Sylvester-Bradley, 1948 Genus Actinocythereis Puri, 1953 Subgenus Ponticocytheres McKenzie, 1967 Actinocythereis (Ponticocythereis) militarts (Brady) (Fig. 50,P) (AM P36476) Cytherers militaris Brady, p.385, pl.61, fig. 9a-d Cythere clavigera Brady, p.111, pl.23, fig. 7a-d Ponticocythereis militaris (Brady); McKenzie, p.96, pl.3, fig. 4, text-figs 40 and 10c-d Cythere clavigera Brady; Puri and Hulings, p.270, pl.16, figs 1,2, text-fig. 4 Ponticocythereis militaris (Brady); McKenzie and Pickett, p.239, fig. 5B Actinocythereis (Ponticocytherers) wchthyoderma Brady (Fig. 6E,F) (AM P36483) Cythere ichthyoderma Brady, p.503, pl.2, figs 22,23 Ponticocythereis quadriserialis Brady; McKenzie, p.98, pl.2, fig. 13 Genus Australimoosella Hartmann, 1978 Australimoosella lauta (Brady) Cythere lauta Brady, p.88, pl.21, fig. 4a-d Cythere lauta Brady; Puri and Hulings, p.280, pl.4, figs 5-8 Genus Hiltermannicythere Bassiouni, 1970 Fliltermannicythere’ basstounn Hartmann (Fig. 5J-L) (AM P36488) Filtermannicythere bassiounit Hartmann, p.91, pl.7, figs 6-14 FAiiltermannicythere basstounit, Hartmann, p.232, pl.5, figs 10-20 FAiiltermannicythere basstounit, Hartmann, p.131, pl.9, figs 8-11, 13-14 PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 168 OSTRACODES FROM PORT HACKING Fig.6. A, B, Echinocythereis melobesioides (Brady, 1880). A, LV external view; B, details of ornamentation. Sample 3, Port Hacking, South West Arm. C, D, Cletocythereis rastromarginata (Brady, 1880). C, LV internal view; D, RV external view. Sample 2, Port Hacking, South West Arm. E, F, Actinocytherets (Ponticocytherets ) ichthyoderma (Brady, 1890). E, LV external view; F, RV internal view. Sample 2, Port Hacking, Gunnamatta Bay. G-I, Pectocythere portjacksonensis (McKenzie, 1967), G, LV internal view; H, RV external view; I, details of ornamentation. Sample 2, Port Hacking, Gunnamatta Bay. J-N, Semicytherura illerti Yassini n.sp. J, LV internal view; K, dorsal view of carapace; L, details of ornamentation, holotype; M, RV external view, holotype; N (same scale as M), ventral view of carapace. Sample 1, Port Hacking, South West Arm. O-S, Cytherura portuswelshpoolensis Hartmann, 1980. O, dorsal view of carapace; P, details of ornamentation; Q, LV external view; R, LV external view; S, RV internal view. Sample 1, Port Hacking, Gunnamatta Bay. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 1880 1976 1880 1967 1979 1984 1880 1976 1981 1982 1880 1974 1981 1981 1880 1967 I. YASSINI AND A. J. WRIGHT 169 Genus Echinocythereis Puri, 1953 Echinocythereis melobesioides (Brady) (Fig. 6A,B) (AM P36485) Cythere melobesioides Brady, p.108, pl.18, fig. la-g Cythere melobesioides Brady; Puri and Hulings, pl.25, figs 1-2 Family HEMICYTHERIDAE Puri, 1953 Genus Cletocythereis Swain, 1963 Cletocythereis rastromarginata (Brady) (Fig. 6C-D) (AM P36482) Cythere rastromarginata Brady, p.83, pl.16, fig. la-d Cletocythereis rastromarginata (Brady); McKenzie, p.95, pl.13, figs 1-2 Cletocythereis cf. rastromarginata (Brady): Hartmann, p.234, pl.6, figs 5-7 Cletocythereis rastromarginata (Brady); McKenzie and Pickett, p.239, fig. 5C-D Genus Quadricythere Hornibrook, 1952 Quadricythere obtusalata (Brady, 1880) Cythere obtusalata Brady, p.91, pl.12, fig. la-c Cythere obtusalata Brady; Puri and Hulings, p.282, pl.5, figs 10-12 Genus Procythereis Skogsberg, 1928 Subgenus Serratocythere Hartmann, 1979 Procythereis (Serratocythere) denswireticulata Hartmann (AM P36500) Procythereis (Serratocythere) densuireticulata Hartmann, p.110, pl.7, figs 3-9 Procythereis (Serratocythere) lyttletonensis Hartmann, p.131, pl.5, figs 6-11 Procythereis (Serratocythere) kerguelenensis (Brady) (AM P36501) Cythere kerguelenensis Brady, p.78-79, pl.4, figs 16-18; pl.20, fig. la-f ‘emicythere’ kerguelenensis (Brady); McKenzie, p.160, pl.1, fig. 9 Procythereis (Serratocythere) australis Hartmann, p.110, pl.7, figs 1-2 Genus Caudites Coryell and Fields, 1937 Caudites litusorienticola Hartmann (Fig. 7H) (AM P36481) Caudites litusorienticola Hartmann, p.111, pl.7, figs 10-13 Family LOXOCONCHIDAE Sars, 1925 Genus Loxoconcha Sars, 1868 Loxoconcha australis Brady (Fig. 7P) (AM P36490) Loxoconcha australis Brady, p.119, pl.28, fig. 3a-d, fig. 5a-f Loxoconcha australis Brady; McKenzie, p.86, pl.12, figs 10-11, fig. 3n-o PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 170 OSTRACODES FROM PORT HACKING Fig. 7. A, B, Paradoxostoma augustensis Hartmann, 1979. A, LV external view; B, RV internal view. Sample 2, Port Hacking, South West Arm. C, D, Paracytheroma sudaustralis (McKenzie, 1978). C, RV external view; D, RV internal view. Sample 2, Port Hacking, Gunnamatta Bay. E, F, Xestoleberis chilensis austrocontinentalis Hartmann, 1978. E, RV external view; F, RV internal view. Sample 1, Port Hacking, Gunnamatta Bay. G, Parakrithella australis McKenzie, 1967. G. RV external view. Sample 1, Port Hacking, Gunnamatta Bay. H, Caudites litusorienticola Hartmann, 1981. H, RV external view. Sample 2, Port Hacking, Gunnamatta Bay. I, J, Xestoleberis cedunaensis Hartmann, 1980. I, RV internal view; J, LV external view. Sample 2, Port Hacking, Gunnamatta Bay. K, L, Leptocythere hartmanni (McKenzie, 1967). K. RV internal view; L, LV external view. Sample 2, Port Hacking, Gunnamatta Bay. M, N, Callistocythere dorsotuberculata paucicostata Yassini and Jones. M, LY internal view; N, LV external view. Sample 2, Port Hacking, Gunnamatta Bay. O, Callistocythere puri McKenzie, 1967. O, RV external view. Sample 2, Port Hacking, Gunnamatta Bay. P, Loxoconcha australis Brady, 1880. P, RV external view. Sample 2, Port Hacking, Gunnamatta Bay. Q, R, Aglaiella dietmarkeysert Hartmann, 1979. Q, RV internal view; R, RV external view. Sample 2, Port Hacking, Gunnamatta Bay. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 I. YASSINI AND A. J. WRIGHT 171 Loxoconcha variolata Brady (Fig. 5Q-S) (AM P36491) 1878 Loxoconcha variolata Brady, p.400, pl.18, fig. 4a-d 1880 Loxoconcha variolata; Brady, p.121, pl.29, fig. 6a-d Genus Loxoconchella Triebel, 1954 Loxoconchella pulchra McKenzie (AM P36492) 1967 Loxoconchella pulchra McKenzie, p.88, fig. 4c 1980 Loxonchella cf. pulchra McKenzie; Hartmann, p.140, pl.12, fig. 1 1984 Loxonchella pulchra McKenzie; McKenzie and Pickett, p.236, fig. 4A-B Genus Hemicytheridea Kingma, 1948 Hemuicytheridea hiltoni Yassini, sp.nov. (Fig. 5A-F) (AM P36486-36487) Derwwation of name. for Mr Neville Hilton, chairman of the Lake Illawarra Management Committee Holotype: one right valve AM P36486 Paratypes: 2 carapaces and one left valve AM P36487 Type Stratum: Recent; sandy mud with Posidonia australis Type Locality: Gunnamatta Bay, Port Hacking estuary Dimensions: Holotype length 225; width 115y Paratype length 205-230y; width 100-115p Description: Carapace small elongate, subreniform; dorsal margin straight. Ventral margin slightly sinuous in anteroventral portion, anterior margin broadly rounded. Posterior end subacute with distinct postero-dorsal cardinal angle. Inner lamella moderately wide at anterior end, narrows at posterior margin. Narrow vestibulum present at both margins. Marginal pore canals straight, about 20 at anterior margin and 11 at the posterior margin. Surface of valve with strongly reticulate ornament, with prominent rib running obliquely from postero-median area to antero-ventral region. Two vertical ridges run- ning from dorsal to ventral margin present at posterior end. Wrinkled structure on surface of reticulation observed at high magnification. Sexual dimorphism present, the male being slightly longer and narrower than the female. Remarks: Hinge structure and muscle scars are characteristic of Hemicytheridea. Vhe species is close to Hemicytheridea crenata (Brady, 1890) but the latter mainly differs by having much more pronounced vertical ridges all over the surface of the valves. Family PECTOCYTHERIDAE Hanai, 1957 Genus Pectocythere Hanai, 1957 Pectocythere portjacksonensis (McKenzie) (Fig. 6G-I) (AM P36498) 1967 ‘Hemicytheridea’ portjacksonensis McKenzie, p.85, fig. 31-}; pl.12, fig. 6 1980 ‘Pectocythere’ portjacksonensis (McKenzie); Hartmann, p.122, pl.5, fig. 17 Fectocythere sp. (Ceduna 120) Hartmann, 1980 (AM P36499) 1980 Pectocythere sp. (Ceduna 120) Hartmann, p.123, pl.3, figs 14-17 PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 172 OSTRACODES FROM PORT HACKING Family CYTHERURIDAE Muller, 1894 Genus Cytherura Sars, 1866 Cytherura portuswelshpoolensis Hartmann (Fig. 60-S) (AM P36484) 1980 Cytherura portuswelshpoolensis Hartmann, p.140, pl.12, figs 2-3 1981 Cytherura portuswelshpoolensis Hartmann, p.121 1984 Cytherurid sp.; McKenzie and Pickett, p.236, fig. 4E Genus Semicytherura Wagner, 1957 Semicytherura illerti Yassini, sp.nov. (Fig. 6J-N) (AM P36502-36503) Derivation of name: for Mr Chris Illert, who provided the samples for this study Holotype: a right valve of a male, AM P36502 Faratypes: two carapaces and one left valve, AM P36503 Type Stratum: Recent; sandy mud with Poszdonza australis Type Locality: South West Arm, Port Hacking estuary Dimensions: holotype length 170; width 182y paratype length 170-180; width 75-85 Description: Carapace medium size, elongate subrectangular, with subdorsal weakly developed caudal process. Dorsal margin straight, ventral margin slight sinuous in the middle. Anterior border well rounded in dorsal view. Carapace is parallel-sided and acuminate at the extremities. Surface of valve with one prominent medio-dorsal longitudinal ridge and two or three medio-ventral ridges. Net- work of intercostal reticulation well developed at the anterior and posterior part of the valves. Intercostal area finely punctate. In ventral view four to five subparallel weakly developed ridges present. Eye tubercle prominent. Inner lamella very wide both anteriorly and posteriorly, curving forms forward strongly in central part of valve. Remarks: Such hinge structures and muscle scars are characteristic of Semuicytherura. Sexual dimorphism is pronounced, the male carapace being much longer and narrower than the female. Family XESTOLEBERIDIDAE Genus Xestoleberis Sars, 1866 Xestoleberis cedunaensis Hartmann (Fig. 71, J) (AM P36504) 1980 Xestoleberis cedunaensis Hartmann, p.149, pl.15, figs 1-4 1984 Xestoleberis cedunaensis Hartmann; McKenzie and Pickett, p.239, text-fig. 5Q-S Xestoleberis chilensis austrocontinentalis Hartmann (Fig. 7E,F) (AM P36505) 1978 Xestoleberis chilensis austrocontinentalis Hartmann, p.128, figs 461-464 1981 Xestoleberis chilensis austrocontinentalis Hartmann, p.122, pl.11, figs 1-12 Family PARADOXOSTOMIDAE Brady and Norman, 1889 Genus Paradoxostoma Fischer, 1855 PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 I. YASSINI AND A. J. WRIGHT 1 Faradoxostoma augustensis Hartmann (Fig. 7A,B) (AM P36496) 1979 Paradoxostoma augustensis Hartmann, p.261, text-figs 188-192 1980 Paradoxostoma augustensis Hartmann, p.157 Family CANDONIDAE Kaufmann, 1900 Genus Aglaiella Daday, 1910 Aglaella dietmarkeyser. Hartmann (Fig. 7Q,R) (AM P36477) 1979 Aglaiella dietmarkeyser: Hartmann, p.264, text-figs 241-250 ACKNOWLEDGEMENTS We thank C. Illert for providing the samples, Mrs Anne Clarke and the Lake Illawarra Management Committee for their support, Therese Carmody who typed the manuscript and David Martin who drafted the figures. The work was supported by a 1986 grant from the University of Wollongong Research Committee. SEM facilities were kindly provided by the Electron Microscope Unit, University of Sydney. References BrRaby, G. S., 1866. — On new or imperfectly known species of marine Ostracoda. Trans zool. Soc. Lond. 5: 359-93. ——., 1878. — A monograph of the Ostracoda of the Antwerp Crag. Trans zool. Soc. Lond. 10: 379-409. ——,, 1880. — Report on the Ostracoda dredged by H.M.S. Challenger during the years 1873-1876. Rept Voyage Challenger, Zool. 1: 1-184. —., 1890. — On Ostracoda collected by H. B. Brady in the South Sea Islands. Trans Roy. Soc. Edinb. 25: 489-524. CHAPMAN, D. M., GEARY, M., Roy, P. S., and THOM, B. G., 1982. — Coastal Evolution and Coastal Erosion in New South Wales. A report prepared for the Coastal Councils of N.S.W. 340pp. CHAPMAN, F., CRESPIN, I., and KEBLE, R. A., 1928. — The Sorrento Bore, Mornington Peninsula, with a description of new or little known fossils. Rec. geol. Surv. Vict. 5: 1-95. Gipps, P. J., 1986. — Five N.S.W. barrier lagoons; their macrobenthic fauna, seagrass communities. Kensington (N.S.W.): University of New South Wales, Ph.D. thesis, unpubl. GODFREY, J. J., and PARSLOW, J., 1976. — Description and Preliminary Theory of Circulation in Port Hacking estuary. C.S\1.R.O. Aust. Div. Fish. Oceanogr. Rept 67. HARTMANN, G., 1978. — Zur Kenntnis des Eulitorals der australischen ktisten unter besonderer Bertick- sichtigung der Polychaeten und Ostracoden. Mitt. hamb. zool. Mus. Inst. 75: 63-219. ——, 1979. — Die Ostracoden der Ordnung Podocopida G. W. Muller, 1894 der warmtemperierten (anti- borealen) West- und Stidwestktiste Australiens (zwischen Perth im Norden und Eucla im Stden). Mitt. hamb. zool. Mus. Inst. 76: 219-301. ——.,, 1980. — Die Ostracoden der Ordnung Podocopida G. W. Miller, 1894 der warm-temperierten und subtropisch-tropischen Ktstenabschnitte der Std- und Stidostkuste Australiens (zwischen Ceduna im Westen und Lakes Entrance im Osten). Mitt. hamb. zool. Mus. Inst. 77: 11-204. ——, 1981. — Die Ostracoden der Ordnung Podocopida G. W. Muller, 1894 der subtropisch-tropischen Ostkuste Australiens (zwischen Eden im Stden und Heron Island im Norden). Mitt. hamb. zool. Mus. Inst. 78: 97-149. ——,, 1982. — Beitrag zur Ostracodenfauna Neueseelands (mit einem Nachtrag zur Ostracodenfauna der Westkuste Australiens. Mutt. hamb. zool. Mus. Inst. 79: 119-50. MCKENZIE, K. G., 1967. — Recent Ostracoda from Port Philip Bay, Victoria. Proc. Roy. Soc. Vict. 80: 61-106. ——, 1974. — Cenozoic Ostracoda of southeastern Australia with the description of Hanaiceratina new genus. Geoscience and Man 6: 153-82. ——, 1978. — Biogeographic patterns in Australian Cainozoic Ostracoda, with the description of Orlo- vibairdia new genus. J. palaeont. Soc. India 20: 279-88. ——,, 1981. — Chapman’s ‘Mallee Bores’ and ‘Sorrento Bore’ Ostracoda in the National Museum of Victoria, with the description of Maddocksella new genus. Proc. Roy. Soc. Vict. 93: 105-7. PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 174 OSTRACODES FROM PORT HACKING ——, 1986. — A comparative study of collections from the S.W. Pacific (Saipan to Tonga), with the descrip- tion of Gambiella caudata Brady, 1890) and new species of Pterobairdia (Ostracoda). J. Micropaleont. 5: 91-108. , and PICKETT, J. W., 1984. — Environmental interpretations of Late Pleistocene ostracode assem- blages from the Richmond River valley, New South Wales. Proc. Roy. Soc. Vict. 96: 227- 42. NEWELL, B. S., 1966. — Seasonal changes in hydrological and biological environments of Port Hacking, Sydney. Aust. J. mar. freshw. Res. 17: 77-91. PurRI, H. S., and HuLINGs, N. C., 1976. — Description of lectotypes of some ostracodes from the Challenger Expedition. Bull. Brit. Mus. (Nat. Hist.), Zoology 29: 251-315. ROCHFORD, D. J., 1951. — Studies in Australian estuarine hydrology. I — Introduction and comparative features. Aust. J. mar. freshw. Res. 2: 1-116. ——., 1959. — Classification of Australian estuarine systems. Arch. Oceanogr. Limnol. Vol. II Supp I. Scott, B. D., 1978. — Nutrient cycling and primary production in Port Hacking, New South Wales. Aust. /. mar. fresh. Res. 29: 803-15. STATE POLLUTION CONTROL COMMISSION (N.S.W.), 1983. — Environmental Audit of Lake Macquarie, 138 p. YASSINI, I., and JONES, B. G., 1987 . — Ostracoda in Lake Illawarra: environmental factors, assemblages and systematics. Aust. J. mar. freshw. Res. 38: 795-843. APPENDIX LIST OF SPECIMENS 36476 Actinocytherers (Ponticocytherets ) militaris 36477 Aglaiella dietmarkeysert 36478 Bairdoppilata sp. 36479 Callistocythere dorsotuberculata paucicostata 36480 Callistocythere puri 36481 Caudites litusortenticola 36482 Cletocythereis rastromarginata 36483 Actinocythereis (Ponticocythereis) ihthyoderma 36484 Cytherura portuswelshpoolensis 36485 Echinocythereis melobesroides 36486 Hemicytheridea hilton: (holotype) 36487 H. hiltoni (paratypes) 36488 ‘Hiltermannicythere’ bassiouni 36489 Leptocythere hartmanni 36490 Loxoconcha australis 36491 L. variolata 36492 Loxoconchella pulchra 36493 Maicrocytherura aestuaricola 36494 Osticythere baragwanathi 36495 Paracytheroma sudaustralis 36496 Paradoxostoma augustensis 36497 Parakrithella australis 36498 — Pectocythere portjackonensis 36499 Psp. (Ceduna 120) 36500 Procythereis (Serratocythere) densuireticulata 36501 P(S.) kerguelenensis 36502 Semicytherura illerti (holotype) 36503 S. illerti (paratypes) 36504 Xestoleberis cedunaensis 36505 X. chilensts austrocontinentalis PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A new Species of Nitella (Characeae) belonging to the Pluricellulate Species Group in Australia* A. T. HOTCHKISS and K. IMAHORI Horcnukiss, A. T., & IMAHORI, K. A new species of Netella (Characeae) belonging to the pluricellulate species group in Australia. Proc. Linn. Soc. N.S.W. 110 (2), (1987) 1988: 175-185. A new species of Nitella, Nitella woodi, belonging to the pluricellulate species group in Australia, is described from the Nepean River, near Camden, New South Wales. The new species appears to be related to Netella cristata and to Nitella hookeri, species endemic to the Australasian region. A. T: Hotchkiss, Professor Emeritus, Department of Biology, University of Louisville, Louisville, Kentucky, 40292, U.S.A., and K. Imahori, Vice-President, Naruto University of Teacher Edu- cation, Takashima, Naruto-Shi, 772, Japan; manuscript received 17 September 1986, accepted for publication 19 August 1987. INTRODUCTION During the course of systematic observations of charophytes in the Nepean River near Camden, New South Wales through the year 1968, a robust form of Nitella was found which 1s described here as a new species, Nitella woodi. The first specimen, taken in March while dredging for Chara australis in a deep pool near the highway bridge north of Camden, was a sterile fragment of a large plant resembling Netella flexilis. In April, studies were concentrated on abundant charophyte beds in the more accessible shallow pools above and below the highway bridge of the airport road west of Camden. A single bed of the new species was apparently restricted to the shaded cooler depths of a pool directly under the bridge. This is the type locality. While the plants in this bed remained sterile and without new growth for several months, all other nearby species, grew and fruited abundantly in beds scattered up and down the river. These included: Chara australis, Nutella sondert, Nutella pentcillata, Nitella tasmanica, Nitella imahorit and Nitella cristata, among dioecious species and Chara gymnopitys, Nutella imperialis and Nitella hortkawi, among the monoecious forms. Meanwhile, in the new species, peculiar vegetative, bud-like growths appeared and were studied from April until August. The growths showed possible dormancy and vegetative reproductive traits suggesting the designation of winter buds or turions. In early spring (late August), a burst of bright green new growth rooted in the bottom mud replaced the darkened, decaying older plants. New turions appeared on the new shoots and fruiting of male and female gametangia was abundant on separated plants from October into November. The late winter early spring appearance of this species suggests that it is markedly seasonal. The bed of the Nepean River presents generally favourable substrate conditions for the growth of charophytes. Coming from a sandstone region, the river winds in the Camden area in a shallow sandy trough over sandbars mixed with varying degrees of organic muck. Periodically flooded, scoured and reshaped by water currents, the sandy bottom presents a dynamic substrate which charophytes are well able to exploit quickly. In 1968, conditions were optimal for the growth of charophytes. The recently-scoured river bed presented broad expanses of clean, sandy bottom. There was a good supply of * A paper presented at the International Seaweed Symposium, Santa Barbara, California, U.S.A., August 1977. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 176 A NEW SPECIES OF NITELLA clean water in a shallow steady flow over rather broad shallow sandbars secure from competition from aquatic angiosperms along the shore. This favourable state continued through the following autumn, winter and into the spring when conditions slowly deteriorated partly as a result of encroaching sedges, silt and epiphytes, but mostly the result of the steadily lowering water level which left stretches of the river semistagnant and eventually in the grip of a drought, but not before the charophytes had produced an abundant crop of spores. In 1976 and in 1980, the first author revisited the site to find it under drought conditions. No collections could be made at either time. OBSERVATIONS Collections: Deep shaded pool, soft muddy bottom, under bridge, airport road, Nepean River, west of Camden, New South Wales. A. T. Hotchkiss 68-3-13-2; 68-3-29-1 (observed in April); 68-6-29-2 (observed July, August); 68-10-5-1; 68-10-19-1; 68-11-3-1 (HOLOTYPE). Specimen, currently in University of Louisville Herbarium, will be transferred to the University of Sydney for disposition. Drawings made by K. Imahori from 68-10-19-1. Figs 1-13. Diagnosis: Nitella woodu A. T’ Hotchkiss et K. Imahori. Figs 1-13 Notella woodii sp. noy. A. T. Hotchkiss et K. Imahori. Planta dioecia, 30-100cm alta, viridis ad brunnea, gymnocarpa. Caulis robustus, 600-1250 crassus; internodiis quam ramuli 2-plo longiora vel aequilonga. Verticilli steriles majores, ramulis 6, 2-3-tim furcatis, 8-11cm longi; radiis primarii totius longitudinis ramulorum 1/3- 2/5, 2.5-4cm longi; radii secundarii 2-4, saepe cum ramulis accessoribus; radiis tertii 2-3; quaterni 2-3, abbreviati. Dactyli 2-3, (1-) 2-4 cellulati, inaequilongi; cellula ultima 55-100y longa, ad basin 25-40p lata. Rami fertiles spiciformes ex apice vel axillari ramulorum secundari. Verticilli feminis quam masculi minus congesti vel similes; ramuli 6, 2-3 (-4) furcatis, 2-5.5cm longi; radii primarii totius longitudinis ramulorum 1/2-3/5; secundarii 3-4; tertii 3-4; quaterni 2-3, quorum 1 saepe in radiis quintis furcati. Dactyli 2-4, 2-3 cellulati; cellula ultima mucronata. Gametangia ad nodos omnes exceptis primariis, solitaria. Antheridia 450p longa et 420 diametro. Oogonia brevi-stipitata, 675 longa, 515y lata; cellulae spirales 7-8; coronula parva, 35 alta, cellula superiore quam inferiori 2-3 tim longiore, ad basin 50-55p lata. Oospora fulva ad purpureo-brunnea, 515 longa et 325y lata; fossa 80p lata; striis 5, paulum prominentibus. Membrana externa moniliformae; medius reticulata, trans ca 32 maculae; interior plane levis. Plant Description: Plants large, usually about 33cm but up to 1m tall; new growth bright green colour, contrasting with older portions darker to brownish green to black; flexible and resembling N. flexzls in the field; without mucus (Figs 11-12). Axes moderately stout to stout in basal internodes, 600-1250 in diameter, not encrusted; internodes about equalling upper branchlets in length to about twice the length of lower branchlets, 6-12cm long. Branches 1 large branch per node together with 1-2 later-formed and apparently secondary, smaller branches from its base; also usually an adventitious branch at the first furcation of a few or all sterile branchlets per whorl and in lower whorls of fertile branchlets. Sterile branchlets 6, 2-3 furcate, up to 8-11cm long, strongly ascending at the upper nodes to a little divergent below (Fig. 7). Primary rays about 0.3 to less than half the branchlet length, 2.5-4cm long. Secondary rays 2-4, one abaxial, slightly PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 177 broader and often monopodial or nearly so, some remain simple; and adventitious branch at first furcation often. Tértiarzes 2-3, divergent but abaxial one less so and nearly or quite monopodial, some remain simple, as long dactyls. Quaternaries 2-3, usually as brachydactyls; one may be monopodial and longer, the other shorter and lateral, divergent. Dactyls 2-3, (1-)2-4-celled, variable in length, usually very long (2-3cm) as secondaries, or shorter (lcm) as tertiary rays, but brachydactylous (.2-.5mm long) as quaternaries (Fig. 5). A typical long dactyl with a long allantoid basal cell abruptly narrowing to a node, a shorter and narrower second cell also abruptly narrowing to a node, a short, conical, mucronate apical cell, or sometimes two cells in the mucro with the penultimate cell longer or shorter and either tapering smoothly into the end cell or the end cell abruptly mucronate. End cell 55-100u long, 25-40 broad at base, (1-celled dactyl: 80u long, 25u broad), often deciduous but leaving a truncate scar. Longer dactyls appear to result from a failure to fork at an upper node. Brachydactyls 2-3, (1-)2-3(-4)- celled, sometimes forming a mucronate 2-3-celled endpiece, smoothly or abruptly tapering down to an acute conical end cell, end cells similar to those on longer dactyls. Fertile branchlets: (Figs 6, 8, 9), male and female gametangia on separate plants in terminal inflorescences consisting of whorls of 6 branchlets at first compacted into closer heads above, later elongating, spreading and widely spaced below (internodes 2.5-8cm) together with fertile axillary branches. The first fruiting appears at the base of an inflorescence with the development of a fertile axillary branch in a whorl of sterile or nearly sterile branchlets. Later there may be a second, smaller branch. A fertile adventi- tious branch may appear at the first furcation of the otherwise sterile branchlets in the basal whorl(s). Upwards in the inflorescence second and third branchlet whorls become progressively more fertile, shorter and accompanied by conspicuous fertile axillary branches. These are followed by several (3-4) whorls of completely fertile branchlets which may or may not terminate the stem. It appears in some cases that the growth of the stem axis continues with the production of further whorls of sterile branchlets above the fertile head. Female inflorescence: lower branchlet whorls 6, 2-3-furcate, from 3-5.5cm long, primary rays 2-2.5cm long, mostly sterile but with fertile axillary branches and occasional adventitious branches at the first branchlet furcation; upper branchlet whorls 6, 2-3(-4)- furcate, form 2-2.5cm long; primary rays 1-1.5cm long; secondaries 3-4, one secondary ray usually broader, less divergent to monopodial; tertiaries 3-4, one may be central; quaternaries 2-3, one may be central and longer; quinaries 2-3, 2-3-celled, tiny, one ray may be longer. Dactyls 2-4, 2-3-celled, 0.3-4mm long in expanded mature branchlets, end cell short conical, mucronate, 50-120u long, 25-35 broad, or a 2-celled terminal mucro, occasionally deciduous. Oogonia: (Fig. 2), short-stalked, solitary; 675 (excl. coronula) long, 515 broad; convolutions 7-8; coronula 35 high, 50-55u broad at base, upper cells overarching, 2-3 times longer than lower. Oospores: (Fig. 1), dark, chestnut brown, 515 long, 325 broad; striae of 5 prominent and flanged ridges; fossa 80m across; membranes: (Figs 3-4), outer, densely granulate to vermiferous; middle, finely reticulate about 32 meshes across; inner, smooth and clear. Male inflorescence: lower branchlet whorls 6, 2-3-furcate, about 3-5cm long, similar to the female in arrangement, composition, and fertility but somewhat more spreading to reflexed, somewhat protandrous. Primaries about half the branchlet length, secondaries 3- 4, tertiartes 2-4, quaternaries 2-3 where present, quinaries 2-3, 2-celled dactyls here present. Dactyls: 2-3, (1-)2-3-celled, up to 1.5cm long, or brachydactylous and (1-)2-3 very short cells in length. Upper branchlet whorls 6, 2-3(-4)-furcate, closer, shorter and more fertile than the branchlets below as in the female. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 178 A NEW SPECIES OF NITELLA Antheridia: solitary at 2nd, 3rd and 4th furcations, absent at the first branchlet node, sometimes terminal at end of tertiary ray or 8 accompanied by minute quaternary dactyls; 450 long, 420 broad, 8-scutate. Chromosomes: A chromosome number ofn = 9 was established for Nitella wood (A. and D. Hotchkiss) unpublished. Turions: Muenscher (1944) calls a turion ‘a hardened abbreviated axis or winter bud as in Potamogeton. The use of the term ‘turior’ herein comes from a combination of suggested morphological and physiological similarity. Winterbud-like structures or turions were observed from March to July on sterile overwintering plants in the Nepean River (Figs 10, 11, 13). The turions consisted of greatly swollen, starch-filled, food-storage cells. Turions in a main stem axis might include a single condensed whorl of primary rays, or primaries, secondaries and some- times tertiaries as well, or two successive branchlet whorls including the connecting internodal cell. Often, a short axillary branch including the stem axis and one of two whorls of branchlets forms an axillary turion. The swollen ray cells form long narrow cylinders gradually tapered at the base, but abruptly tapered at the apex, and all become quite rigid. Turions were brought into the Jaboratory and cultured in pans of water under lights in a constant temperature room. Germination was first observed in June after 4 to 6 weeks of cultivation. New growth from the turions produced normal stems and branchlets. Occasionally in sterile branchlet whorls, more rarely in fertile whorls, new turions appear in the upper shoots with the new growth of early spring (August) and production of turions continues through the fruiting season (October-November). At this time it was seen that young turions may be fertile on normal nodes at a few furcations above the swollen turion rays. An attempt to germinate new turions was unsuccessful through December when the experiment was terminated. Turion cells appear to be longer-lived and more resistant to adverse conditions than are the ordinary vegetative cells in both culture and in the field where they often become detached as a group and survive when other nearby cells are dead. It is likely that detached turions can serve as propagules and disseminules for the species. It seems possible that young turions possess some degree of dormancy which may assist in this role. Turions such as these are apparently a useful taxonomic tool for delimiting Netella woodu from many charophytes. They are probably not restricted to this species. For example, the second author found ‘turions’ in Japanese specimens of Nitella annularts. In addition, Wood and Imahori (1965) provided illustrations of Nitella tumida and Chara sub- mollusca, which strongly suggest the presence of turions in the enlarged, inflated primary rays in certain whorls of branchlets. RELATIONSHIPS The establishment of this member of the genus Nitella as a new species was the result of a series of studies and observations. (1) This large, robust species, complete with its unusual turions, was unique among all previously-studied forms. (2) The differences in morphology coupled with its seasonal maturation in late winter, early spring, distinguished it from other charophytes in the Nepean River area. (3) In its strong tendency to monopodial branchlet axis rays, the new species might be considered close to the Nutella cristata complex, (Wood and Imahori, 1965). It 1s PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 179 easily separated from N. cristata by additional morphological characteristics and by its difference, locally, in maturation time. (4) As in much of the genus Nitella, the relationships of this species are most clearly indicated by the nature and number of cells in the dactyls. Its 1-4-celled dactyls place the species in the pluricellulate series of the Arthrodactylae. A combination of morphological characters narrows it down within the confines of Wood’s key (1965) to the Section Incertae which contains Nitella hooker, tentatively assigned as a form of the subgenus Netella (which also includes Nitella flexilis) and Nitella tasmanica, in the subgenus HZyella. The species resembles dioecious N. tasmanica, somewhat, but in general habit and in the form of its dactyls, it is closer to the monoecious N. hookeri. The two may be separated by N. hooker’s geminate oogonia and coarsely reticulated oospore membrane. A chromosome number of n = 18 (A. and D. Hotchkiss, unpublished) was established for the monoecious N. hooker: (sensu N. tricellularis, fide R. D. Wood, collected by Wood in New Zealand) in 1961, whereas the dioecious Nitella woodii has a chromosome number ofn = 9. It would appear that the morphology of the species, its dioecious condition and its chromosome number provide sufficient distinctions to establish Nitella woodi as a separate species. The new species has been named Nitella wood in honour of the late Professor R. D. Wood, a leading student of the Charophyta. ACKNOWLEDGEMENTS We wish to express our deepest appreciation for the interest shown and the help extended in collection and preservation of materials by the late John Waterhouse of the University of New South Wales. References MUENSCHER, W. D., 1944. — Aquatic Plants of the United States. (x + 374). Ithaca, N.Y.: Comstock Pub. Co. Woop, R. D., and IMAHORI, K., 1965. — A Revision of the Characeae. Vol. 1. Monograph of the Characeae (by R. D. Woop, illustrated by K. IMAHORI). (xxiv + 904). Weinheim, Germany: J. Cramer PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 180 A NEW SPECIES OF NITELLA Figs 1-5. 1, Oospore, x90; 2, Oogonium, x75. 3, Spore Membrane, lower focus, x600. 4, Spore Membrane, higher focus, x600. 5, Dac tyls at apices of sterile branchlets, x75. PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 181 Fig. 6. Fertile female branchlet, x30. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 182 A NEW SPECIES OF NITELLA Figs 7-9. 7, Sterile branchlet, x4.5. 8, Male branchlet, x9. 9, Female branchlet, x6. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 183 Fig. 10. Turions in two whorls of branchlets, x7.5. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 184 A NEW SPECIES OF NITELLA ———— ——— | 12 Figs 11-12. 11, Habit of female plant, turions in lower branchlet whorl, x3/2. 12, Habit of male plant, x3/2. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 185 Fig. 13. Turions stained with iodine. Slide prepared and photographed at the University of Sydney. PROC. LINN. SOG. N.S.W., 110 (2), (1987) 1988 Additional Observations on Nitella verticillata (Characeae) from anew Locality in New South Wales* A. T. HOTCHKISS and K. IMAHORI HorcnHkiss, A. T., & IMAHORI, K. Additional observations on Nitella verticillata (Characeae) from a new locality in New South Wales. Proc. Linn. Soc. N.S.W. 110 (2), (1987) 1988: 187-191. Abundant fertile material of dioecious Netella verticillata (with mucus) from brack- ish coastal lagoon Lake Munmorah, New South Wales, permits description of’ monopodial male plants, antheridia, oogonia and spores, in a species heretofore known only from limited material in Western Australia, South Australia and Victoria. A. T. Hotchkiss, Professor Emeritus, Department of Biology, University of Louisville, Louisville, Kentucky, 40292, U.S.A., and K. Imahori, Vice-President, Naruto University of Teacher Edu- cation, Takashima, Naruto-Shi, 772, Japan; manuscript received 17 September 1986, accepted for publication 19 August 1987. INTRODUCTION Nitella verticillata was first collected by Nancy Burbidge (1956, 1960), from Lake Parkeyerring, a shallow salt water lake near Wagin, Western Australia, June 5, 1933. The specimens she sent to G. O. Allen in England proved to be sterile, incomplete, and scanty. Allen sent the material to Filarszky (1937) in Hungary, who in turn described it and set it up as a monotypic genus, Charina, and established Charina verticillata. Although no gametangia were available the new species was designated monoecious. Wood and Imahori (1964, 1965) further described and illustrated the species from the incomplete sterile specimens available from G. O. Allen, and added some details. The species was transferred at this time to the genus Nitella by Wood. Wood (1972) reported finding the species in two new localities: Lake Bunijon, Victoria, and the Thorndon Park Reservoir near Adelaide, South Australia. He pointed out that the material was ‘so small and threadlike’ that during the course of a year collect- ing in Australia, he did not detect it in the field, but found it only because fragments were entangled with plants of other Characeae collected in Victoria and South Aus- tralia. The material yielded female gametangia and oospores which he described and led him to suggest that the species might be dioecious rather that monoecious. He summarized the habitat of the species in the three Australian States as follows: ‘shallow water of flooded field or lake; firm bottom; fresh water, possibly tolerant of at least slight salinity. Finally, Wood separated the new material in his key on the basis of its bicellu- late dactyls, its monopodial branchlets and its small size. In February, 1976, the first author collected Noetella verticillata at Lake Munmorah, north of Sydney, New South Wales. Lake Munmorah is a coastal lagoon lake with an inlet to the sea opening between sand dunes along the coast. Salinity varies with the rainfall. At the time of collecting, after an unusually rainy season with high water, the salinity level was reported to be 12.3 p.p.m. In time of drought, this varies up to 18 p.p.m. The species was found growing with Lamprothamnium and Chara species. Nitella verticillata * A paper presented at the International Seaweed Symposium, Santa Barbara, California, U.S.A., August LOWE PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 188 OBSERVATIONS ON NITELLA VERTICILLATA PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 189 was growing on the flooded sandy, pebbly beaches with Lamprothamnium and Chara in deeper water nearby. OBSERVATIONS Collections: Lake Munmorah near Ashdam Creek and power plant inlet canal. A. T. Hotchkiss 76-2-24-1. Drawings made by K. Imahor, Figs 1-14. Description: Nitella verticillata (Fil. et G. O. Allen ex Fil.) R. D. Wood Plants dioecious, male and female plants similar vegetatively, bright to dark green, much branched and growing together, up to 8cm high, (Fig. 1). Axis moderately slender to 330u in diameter: internodes as long as fertile whorl, 2-3 times as long as sterile branchlets. Branchlets 6-8 in a whorl, 1-3 furcated, (Fig. 5). Fertile branchlets covered with mucus and commonly congested into heads to 2mm long; primary rays 4 to % of the entire length; secondary rays 3-4, of which 1 is usually the central ray, but absent when an oogonium or antheridium is produced terminally, and 1-3 lateral rays being dactyls usually; tertiary rays 4-6 (very rarely only 1) of which central and some lateral rays fur- cate once more into 4-6 terminals. Branchlet nodes on male plants commonly whorls of six lateral rays plus short central axis terminating in an antheridium which 1s solitary and octoscutate. Sterile branchlets somewhat diffuse (Fig. 8), 1-2 times furcated; primary rays */3 to °/s of the entire length; secondary rays 4-6 of which 1 is occasionally central; tertiary rays 3-4. Dactyls (Figs 4, 6), elongated on fertile branchlets and at the primary node of the sterile branchlets; abbreviated and percurrent usually at the primary node of the sterile branchlets, abbreviated and percurrent usually at the second node of fertile branchlets; uniformly 2-celled but seems unicellular because of the deciduous end cells, (Fig. 6); end cells acute or acuminate, 39-100p wide at the base (Fig. lal) Occasionally 1 to 4 dwarf branchlets or accessories produced at the base of sterile whorls, (Figs 12-14), 200 to 600u long; branchlets simple (as a single dactyl) or once furcated into 3-4 dactyls. Gametangia produced at the Ist and 2nd nodes of branchlets. Oogonia solitary or geminate, (Fig. 2), terminal or laterally produced; 550 to 580 long (including the coronula) and 375 to 400u wide; spiral convolutions 8-9; swollen at the base of coronula, (Fig. 7). Coronula small, 35-40pn high, 60 to 65 wide at the base. Oospores bright to dark brown (Fig. 9); subglobose and compressed, 320 to 350u long, 280 to 300 wide and 180 to 200 thick; striae prominent and flanged, 6-7; fossa 50 to 554; membranes (Fig. 10), bright brown, finely reticulated; 12 to 14 meshes across fossa, arranged in a line series. Antheridia (Fig. 3), solitary and terminal; stalk cell 30 to 60p long by 30p diameter; antheridia 250 to 330u diameter. Chromosome number, n = 9. Figs 1-11. 1, Plant habit, with mucus around heads, natural size. 2, Geminate, lateral oogonia, with secondary rays surrounding central monopodial, secondary ray, x35. 3, Solitary antheridium, terminal on short, central, secondary ray, x35. 4, Short, ovoid dactyls, x35. 5, Part of a female plant showing whorl of branchlets, x15. 6, Dactyls from which the end cells have fallen off, x35. 7, Apex of oogonium, somewhat swollen at the base of coronula, x150. 8, Part of a sterile whorl of 6 branchlets, x150. 9, Oospore, with 6-7 prominent and flanged striae, x65. 10, Oospore membrane, finely reticulated, 12 to 14 meshes across fossa arranged in a somewhat linear series, x270. 11, End cells of dactyls mucronate, acute or acuminate, often deciduous, x150. (Linear scale for Figs 1-11 is 20 mm.) PROG. LINN. SOG. N.S.W., 110 (2), (1987) 1988 190 OBSERVATIONS ON NITELLA VERTICILLATA Figs 12-14. Dwarf accessory branchlets, x35. 12, Non-furcating branchlet. 13, Non-furcating and once furcated branchlets. 14, Two dwarf branchlets produced between central and lateral rays. (Linear scale for Figs 12-14 is 20 mm.) RELATIONSHIPS Wood (1964, in Wood and Imahori, 1964), without benefit of gametangia, placed Nitella verticillata in his subgenus Tveffallenia, Section 8, Migularia with three other species, Nitella struthioptila, N. imahori and N. cristata. The positioning was based strictly on the arrangement of the percurrent central branchlet ray, a type of branchlet found in a few other species including the N. cristata complex. The small N. vertzcillata with its 1-2 cell dactyls is easily separated from the larger N. cristata with its 3-4 cell dactyls. A new species described by Williams (1959), Nitella reticulata from a coastal sand dune pond near Sydney, with its reticulate spore membrane and brackish water habitat, PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. T. HOTCHKISS AND K. IMAHORI 191 is suggestive of N. verticillata but its dactyls, furcation and size serve to separate it from N. verticillata. The Nitella hyalina species group, in the same subgenus Tveffallenia, but in Section 4, Decandollea, is often found in saline water or near coastal areas. The group has strongly percurrent branchlet axes and an abundance of mucus. However, its well-developed accessory branchlets and its much larger size separate it easily from WN. verticillata. The additional material provides an opportunity for further study of the position of the relatively obscure Netella verticillata within the genus Nitel/a and of the importance of several key characteristics, such as the percurrent central branchlet ray. More extensive data on oogonial structure and the oospore membrane will fix its relationships more fully. In additon, the new details on antheridial structure and the arrangement of the male branchlets, enables this species to be subject to as critical analysis and comparison as any of the best known members of the Characeae. References BURBIDGE, N. T., 1956. — Robert Brown’s Australian collecting localities. Proc. Linn. Soc. N.S.W. 80: 229-233. —, 1960. — The phytogeography of the Australian region. Aust. J. Bot. 8: 75-212. FILARSKY, N., 1937. — Idigenfoldi Charafélék hatarozasa. (Determinatio Characearum exoticarum). Magyar Tud. Akad. Mat. Term. Ert. 55: 476-497, 6 pl. incl. 46 fig. WILLIAMS, M. B., 1959. — A revision of Nitella cristata Braun (Characeae) and its allies. Part II. Taxonomy. Proc. Linn. Soc. N.S.W. 84: 346-356. Woob, R. D., 1972. — Characeae of Australia. Nova Hedwigia 22,11:1-120. , and IMAHORI, K., 1964. — Vol. 2. Iconograph of the Characeae. A Revison of the Characeae. Weinheim: Cramer. , and , 1965. — Vol. 1. Monograph of the Characeae. A Revision of the Characeae. Weinheim: Cramer. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A new Australian larval Callidosomatine Mite (Acarina: Erythraeidae) parasitic on Flies, with Notes on Subfamily and ‘Tribe Classification RV, SOULHCOR, SOUTHCOTT, R. V. A new Australian larval callidosomatine mite (Acarina: Erythraei- dae) parasitic on flies, with notes on subfamily and tribe classification. Proc. Linn. Soc. N.S.W. 110 (2), (1987) 1988: 193-204. A larval mite Carastrum ferrari n. gen., n. sp. (Acarina: Erythraeidae: Callido- somatinae: Callidosomatini), parasitic on Diptera is described from New South Wales. The larvae, generally one per fly, select the ventral surface of the host fly for attachment, in most cases the metasternal-abdominal tergite I membrane. A revised key to the genera of larval Callidosomatini is given, and formal definitions of the tribal names Callidosomatini and Charletoniini. R. V. Southcott, 2 Taylors Road, Mitcham, Australia 5062; manuscript received 24 February, 1987, accepted for publication 25 August 1987. INTRODUCTION The subfamily Callidosomatinae was introduced by Southcott (1957), and later (Southcott, 1961b) divided into the tribes Callidosomatini and Charletoniini, in keys, but hitherto without formal text definitions for the tribes. Southcott (1972) left the Callidosomatini with three genera known as larvae: Caeculisoma Berlese, 1888, Callidosoma Womersley, 1936 and Momorangia Southcott, 1972. In this paper a further genus of Callidosomatini is described, from larvae captured ectoparasitic on flies in New South Wales, Carastrum ferrari n. gen., n. sp. Abbreviations for scutal dimensions are as in Southcott (1966, 1972), and for leg segmental dimensions as in Southcott (1986). Seta coding terminology follows Southcott (1961b,c, 1966, 1972). All measurements are in micrometres (um) unless otherwise indicated. SYSTEMATICS Tribe CALLIDOSOMATINI Southcott Callidosominae Southcott, 1957, p. 97. Callidosomatinae Southcott, 1961b, p. 521. Callidosomini Southcott, 1961b, p. 522. Callidosomatini Southcott, 1972, p. 1. Diagnosis Adults: Callidosomatinae with usually distal tubercles on pedal tibiae. Larvae: Callidosomatinae with scutalae 3+3 or more. Pedocoxalae 1, 2, 2. Pedotrochanteralae 1, 1, 1. Posterior pedotibial claw hooklike. Palpal tibial claw (odontus) without accessory peg or basal process. Type genus Callidosoma Womersley, 1936. Comment: The tribe Callidosomatini was based primarily on the characters of the adults (and deutonymphs), these being generally mites with short, thick legs with prominent distal tubercles. Since then Sharma, Farrier and Drooz (1983) have described the adult and deutonymphal instars of Callidosoma metzi Sharma, Drooz and Treat, 1983 from North PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 194 CALLIDOSOMATINE MITES Q -)) SS) Fig. 1. Carastrum ferrari n. gen., n. sp. A, Dorsal view of Holotype larva, partly in transparency, with legs omitted beyond trochanters. B, Dorsal idiosomal seta (‘ocular’), from Paratype ACA1892B. (Both figures to nearer scale.) PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 R. V. SOUTHCOTT 195 and Central America, and record (both papers) that tibial tubercles are lacking in both of these instars. The larva, however, answers to the criteria of Callidosoma of Southcott (1972). Treat (1985) has recorded that the deutonymphs of Callidosoma apollo Southcott from North America also lack tibial tubercles. He pointed out various overlaps of the characteristics of the larvae of Callidosoma and Caeculisoma, and stated that there are thus unresolved problems in the classification of ‘generic or tribal taxa’. See also the comment after the definition of Charletoniini, below. Key to genera of larval Callidosomatini lame Scutalae4 Fa or mone whee War BE. Wie widle ie eedeng canoes Momorangia Southcott SC UU ee ey hy a es vl rae re Earns A Se es Pa cai cena 2 ee ealpalitilbiall claw wathi4-9 termimaliteeth) . 2 40834. e.cs =e Carastrum n. gen. Palpalstibialsclawi wath) twO) PROMS (5) wae 5 yo we tet ae celts eu ed ce des 3 3. Anterior scutal sensilla not posterior to level of ML scutalae ...... Callidosoma Womersley Anterior scutal sensilla posterior to level of ML scutalae .. Caeculisoma Berlese. Carastrum n. gen. Etymology: The name Carastrum is derived from the Ca- of Callidosomatini, and Latin rastrum, a rake, referring to the structure of the palpal tibial claw. Diagnosis (based upon larval characters): Callidosomatini with scutalae 3+3. Palpal tibial claw with 4-5 terminal teeth. All tarsal claws hooklike, the posterior with three large ventral setules. Pedotibia II with a large solenoidala. Type species Carastrum ferrari n. sp. Carastrum ferrari n. sp. Figs 1 A,B, 2A-C, 3, 4A-H, 5, 6 Etymology: The specific name is in honour of the collector. Material examined (larvae only; adults and deutonymphs not known): Holotype with label ACA1892G2, parasitic on fly A2808, 2 miles [3.2 km] W of Durras (north of Bate- mans Bay), New South Wales, 14.11.1972, P. Ferrar. Paratypes: Nine other specimens, same locality, date and collector, on flies Pyrellia tasmaniae Macquart, Helina sp., and undetermined calyptrates. Mites labelled ACA1892A-F, Gl, H1, H2, on flies A2802-8, 2809, 2809 respectively. All mites deposited in South Australian Museum collection; host Diptera in Australian National Insect Collection, Canberra. Description of Holotype (supplemented by paratypes): Colour red. Ovoid, flattened below, with some grooves of idiosoma extending to lateral outlines. Length of idiosoma in ethanol 585, width 415, overall length including mouthparts 640; after mounting through lactic acid- Hoyer’s medium length 610, width 440, overall length 675. Dorsal scutum rounded, with shallow concave anterior margin, convex lateral margins except immediately before the protrusions for the posterior sensilla, with a notch. between. Anterolateral angles rounded. Scutalae normal, curved, tapering, slightly blunted, with strong setules; AL at AL angles of shield, ML a little behind and lateral to them, PL further lateral but anterior to mid-level of shield. Scutal sensillary setae filiform, tapering, with a few distal setules. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 196 CALLIDOSOMATINE MITES ‘TABLE 1 Standard Data of three specimens of Carastrum ferrari ACA1892G2 ACA1892A ACA1892B Holotype Paratype Paratype AW 61 59 oH MW 66 69 66 PW 71 73 71 SBa 11 10 9 SBp 15 14 14 ASBa 23 25 26 ISD 55 55 56 IL, 85 86 89 WwW 80 90 88 A-M 11 14 15 A-P 31 35 35 AL 55 49 58 ML 55 58 39) PL 52 Aci) 48 ASens 45 45 42 PSens 69 66 67 ASBa/ISD 0.42 0.45 0.46 DS 46-53 45-49 46-55 Oc. 46 = 47 MDS 46 45 515) PDS 53 49 46 Gel 91 87 87 Til 109 112 112 Tal(L) 98 ITS, 109 Tal(H) 19 19 19 Gell 80 77 82 Till 93 102 108 Tall(L) 107 104 107 Tall(H) 19 18 19 Gelll 83 82 87 Tilll 160 155 157 Talll(L) 115 114 120 Talll(H) 19 17 17 The Standard Data of the Holotype and two Paratypes are as in Table 1. Eyes 1+1, posterolateral to scutum, corneae circular, width 18. Dorsum of idiosoma with 28 setae, curved, slightly tapering, blunt-pointed, with strong setules; arranged 2 (oculars), 4, 4, 6, 6, 4, 2. Venter: sternalae I robust, tapering, pointed, well setulose, 45 long; sternalae II similar, 53; sternalae III similar, 53. Behind coxae III 14 similar setae, arranged 4, 4, 4, 2; 29-45 long, the more posterior shorter and tending to resemble posterior dorsal idiosomalae. Coxalae 1, 2, 2; coxala I tapering, pointed, well setulose, 76; lateral coxala II similar, 48, medial coxala II similar, 75; lateral coxala III similar, 50; medial coxala III similar, 60. Legs of normal callidosomatine build. Specialized setae (from ACA1892B): Genu I with SoGel.81d(20 long), VsGel.87pd(5). Tibia I with Cp.61d(6) + SoIil.62d(20) (‘duplex pair’), Solil.69d(27), VsTil.84pd(5). (Most distal seta of tibia I is ScTi1.83v(22) ). Tarsus I with Solal.35d(27), SoFal.51d(2). Genu II with VsGell.87pd(5). Tibia II with Solill.09d(31). Tarsus II with SolaI1.45d(24). Tibia III with greatly enlarged solenoidala Soli111.04d(66). Pretarsal formula 1, 1, 1. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 R. V. SOUTHCOTT 197 30 um ve ioe Fig. 2. Carastrum ferrari n. gen., n. sp. A, Ventral view of Holotype larva, with legs omitted beyond trochanters. B, C, Posterior ventral setae of idiosoma, from Paratype ACA1892B. (All figures to nearest scale.) PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 198 CALLIDOSOMATINE MITES } i) \ 1 Zi/ \ Fig. 3. Carastrum ferrari n. gen., n. sp., larva. Legs I, II, III of Paratype larva ACA1892B, to standard symbols; to scale on left. Inset: Tips of tarsi I and II, to scales alongside: I dorsal aspect, II posterior aspect; Pt pretarsala. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 R. V. SOUTHCOTT 199 PaScTi1 400 um Fig. 4. Carastrum ferrari n. gen., n. sp., Larva. A, Dorsal scutum. B, Gnathosoma, dorsal view. C, Gnathosoma, ventral view. D, Palpal tibial claw, medial aspect, in dorsal view (see B). E, Tip of palp, ventral aspect (see C). (A-E from Paratype ACA1892B.) F, Holotype, dorsal view, in preservative before clearing. G, Large speci- men, Paratype ACA1892C, after mounting, ventral aspect showing gnathosoma on ventral surface. H, Large specimen, dorsal aspect, in preservative, Paratype ACA1892G1, showing dorsal scutum retained in position at anterior pole of idiosoma. (All to nearest scale.) PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 200 CALLIDOSOMATINE MITES Gnathosoma robust, chelicerae rounded, width (combined) 71, length 91. Cheliceral fangs curved, with oblique terminal cutting edge. Gnathosomal lip fim- briated. Anterior hypostomala tapering, pointed, well setulose, 24; posterior similar, 66. Galeala slender, pointed, nude, 27. Palpal setal formula 1, 1, 3, 7. Palpal femorala arises laterally, slender, pointed, setulose, 65; palpal genuala arises dorsally, similar, 40. Other palpal setae as figured. Palpal tibial claw broadened at end, to carry 4-5 small, pointed teeth. Palpal supra- coxala a short blunted peg, 6 long. Remarks on the type series These larval mites were all partly fed. The smallest available, and hence most suitable for detailed description, was selected as the Holotype. Swelling of the idiosoma due to feeding in larger specimens obscures the characters of the gnathosoma. Details of the sizes of the larvae (measured in ethanol, except A. B, which are estimated from the slide mounts) and host identifications are given in Table 2. TABLE 2 Sizes and host Diptera identifications of Carastrum ferrari n. gen., n. sp. H = Helinasp.; PT = Pyrellia tasmaniae Macquart (Muscidae). MF = unnamed calyptrate. Length Width Mite number of of Fly Fly ACA1892 mite mite number identification A ca 1500 ca 900 A2802 H B ca 600 ca 400 A2803 MF C 990 610 A2804 PT D 1585 1045 A2805 MF E 1870 1280 A2806 H F 1530 1115 A2807 H Gi 1600 1175 A2808 H G2 585 415 A2808 H H1 1100 650 A2809 H H2 845 470 A2809 H Remarks on biology All of the mites were attached to the ventral surface of the fly, and all except one to the membrane connecting the metasternum and abdominal tergite I. In the two cases in which two mites attached to the host fly, in one the two mites were attached to the metasternum-abdominal tergite I membrane, and in the other case one mite was on this membrane, and the other on the prosternum. The largest of the larval mites (ACA1892E) equalled in size the abdomen of the host fly (see Fig. 5). Mites of the tribe Callidosomatini have been recorded as larval ectoparasites of Lepidoptera (Womersley, 1934; Southcott, 1972; Treat, 1975, 1985; Sharma, Drooz and Treat, 1983; Sharma, Farrier and Drooz, 1983), Homoptera (Southcott, 1946, 1972) and Orthoptera (Southcott, 1961a, 1972). Although this is the first Australian record of Diptera being parasitized by members of the Callidosomatini, there 1s a previous record of one species, Callidosoma tiki Southcott, parasitizing an unidentified dipteran in New Zealand (Southcott, 1972), and mites referred to Callidosoma spp., but not further deter- mined, have been recorded from Culicidae in North and Central America by Mullen (1975), Welbourn (1983) and Treat (1985), and from Trichoptera in North America by Resh and Haag (1974). PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 9 R. V. SOUTHCOTT 201 TM mh iy Lunn) Fig. 5. Carastrum ferrari n. gen., n. sp., Larva. Mites as single parasites on host Diptera, Paratypes. C, Specimen ACA1892C on its host, Pyrellia tasmaniae Macquart, in lateral aspect. E, E, Lateral and ventral aspects of mite ACA1892E on its host fly (Helina sp.). PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 202 CALLIDOSOMATINE MITES Fig. 6. Carastrum ferrari n. gen.,n. sp., Larva. Host Diptera (Helina sp.) each with two mite ectoparasites. H, H, Lateral and ventral views of Paratypes ACA1892H1, H2 attached to metasternal-abdominal tergite I membrane. G, G, Lateral and ventral aspects of host fly with Holotype ACA1892G2 attached to metasternal- abdominal tergite I membrane, and with Paratype ACA1892G1 attached to prosternum. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 R. V. SOUTHCOTT 203 Tribe CHARLETONIINI Southcott This name was proposed in a key by Southcott (1961b, p. 522), but otherwise has not been formally defined. The following can now be submitted: Definition Adults: Callidosomatinae without distal tubercles on pedal tibiae. Larvae: Callidosomatinae with scutalae 2+2 or 3+3. Pedocoxalae 1, 1, 1 or 2, 2, 2. Pedotrochanteralae 1, 1, 1 or 2, 2, 2. Posterior pedotibial claw hooklike or not. Palpal tibial claw (odontus) with or without accessory peg or basal process. Type genus Charletonia Oudemans, 1910. Comment: Adult Charletoniini can be separated from the Callidosomatini by a single criterion, 1.e. by the absence of distal tubercles on pedal tibiae (see, however, some North American exceptions mentioned earlier). In the larvae so far allotted to the Charletoniini, only one genus has been experimentally correlated with its adult or deutonymphal instar, that being Charletonia, founded by Oudemans on 1 May 1910, and its post-larval instars being described under Sphaerolophus Berlese, founded on 1 July 1910. Correlations have been made by Ishu (in Southcott, 1961b, p. 528) and by Rosa and Flechtmann (1980) and Treat (1980). The other genera founded on larvae, Haupt- mannia Oudemans, 1910, Andrevella Southcott, 1961, Grandjeanella Southcott, 1961 and Pussardia Southcott, 1961, may be tentatively allotted to the Charletoniini on present evidence. All of them come within the definition of Charletoniini as given above, and are excluded from the Callidosomatini as defined earlier in this paper. Further placements of these genera will probably need to depend on the evidence of experimental larva- postlarval instar correlations. ACKNOWLEDGEMENTS I am indebted to Mr Paul Ferrar, formerly of CSIRO Division of Entomology, Canberra, A.C.T., for the specimens, and to Dr D. H. Colless, CSIRO Division of Entomology, for the identifications of the host Diptera. I thank the Australian Biological Resources Study for support. References BERLESE, A., 1888. — Acari Austri-Americani quos collegit Aloysius Balzan. Boll. Soc. Ent. Ital. 20: 171-222. ——,, 1910. — Brevi diagnosi di generi e specie nuovi di Acari. Redia 6 (2): 346-388. MULLEN, G. R., 1975. — Acarine parasites of mosquitoes. I. A critical review of all known records of mosquitoes parasitized by mites. J. med. Entomol. 12 (1): 27-36. OUDEMANS, A. C., 1910. — Acarologische Aanteekeningen XXXII. Ent. Ber., Amst. 3 (53): 67-74. RESH, V. H., and HaaG, K. H., 1974. — New records of parasitism of caddisflies by erythraeid mites. _/. Parasitol. 60 (2): 382-383. Rosa, A. E., and FLECHTMANN, C. H. W., 1980. — Sphaerolophus a synonym of Charletonia? (Acari: Erythraei- dae). Internat. J. Acarol. 6 (3): 215-217. SHARMA, G. D., DRooz, A. T., and TREAT, A. E., 1983. — A new species of Callidosoma (Acari: Prostigma [sic]: Erythraeidae) parasite on adults of Anacamptodes vellivolata (Lepidoptera: Geometridae) as a larva, and predator of eggs of Lepidoptera as deutonymph and adult. Ann. Entomol. Soc. America 76 (1): 120-129. ——, FarRRIER, M. H., and Drooz, A. T., 1983. — Life-history, and sexual differences of Callidosoma metzi Sharma, Drooz and Treat (Acarina: Erythraeidae). Internat. J. Acarol. 9 (3): 149-155. SOUTHCOTT, R. V., 1946. — Studies on Australian Erythraeidae (Acarina). Proc. Linn. Soc. N.S.W. 71 (1-2): 6-48. ——, 1957. — The genus Myrmicotrombium Womersley 1934 (Acarina, Erythraeidae), with remarks on the systematics of the Erythraeoidea and Trombidioidea. Rec. S. Aust. Mus. 13 (1): 91-98. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 204 CALLIDOSOMATINE MITES —, 1961a. — Notes on the genus Caeculisoma (Acarina: Erythraeidae) with comments on the biology of the Erythraeoidea. Trans Roy. Soc. S. Aust. 84: 173-178. ——, 1961b. — Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Aust. J. Zool. 9 (3): 367-610. ——, 1961c. — Description of two new Australian Smarididae (Acarina), with remarks on chaetotaxy and ge- ographical distribution. Trans Roy. Soc. S. Aust. 85: 133-153. ——, 1966. — Revision of the genus Charletonta Oudemans (Acarina: Erythraeidae). Aust. J. Zool. 14 (4): 687-819. ——., 1972. — Revision of the larvae of the tribe Callidosomatini (Acarina: Erythraeidae) with observations on post-larval instars. Aust. J. Zool., Suppl. Ser. 13: 1-84. ——., 1986. — Studies on the taxonomy and biology of the subfamily Trombidiinae (Acarina: Trombidiidae) with a critical revision of the genera. Aust. J. Zool., Suppl. Ser. 123: 1-116. TREAT, A. E., 1975. — Mites of moths and butterflies. Ithaca and London: Comstock Publishing Associates, Cor- nell University Press. ——, 1980. — Nymphal Sphaerolophus reared from larval Charletonia (Acarina: Erythraeidae). Internat. J. Acarol. 6 (3): 205-214. ——, 1985. — Larval Callidosoma (Acari, Erythraeidae): new records and a new rearing from the Americas. Internat. J. Acarol. 11 (2): 93-124. WELBOURN, W. C., — 1983. — Potential use of trombidioid and erythraeoid mites as biological control agents of insect pests, pp. 103-140, zn Hoy, M. A., CUNNINGHAM, G. L., and KNUTSON, L., (eds), Biological control of pests by mites. Berkeley: Univ. Calif., Spec. Publicn. 3304. WOMERSLEY, H., 1934. — A revision of the trombid and erythraeid mites of Australia with descriptions of new genera and species. Rec. S. Aust. Mus. 5 (2): 179-254. ——.,, 1936. — Additions to the trombidiid and erythraeid acarine fauna of Australia and New Zealand. /. Linn. Soc. Lond. (Zool.) 40 (269): 107-121. PROC. LINN. SOC. N.SW., 110 (2), (1987) 1988 The Lichen Genus Cladonia Section Cocciferae in Australia A. W. ARCHER ARCHER, A. W. The lichen genus Cladonza section Cocciferae in Australia. Proc. Linn. Soc. N.S.W. 110 (2), (1987) 1988: 205-213. Eight species of Cladonia Hill ex Browne sect. Cocciferae (Delise) Dahl occur in Aus- tralia. C. angustata Ny]. is reported for the first time in Australia. A key to the species is presented and the major phenolic compounds of each taxon are reported. A. W. Archer, Division of Analytical Laboratories, Department of Health, PO. Box 162, Lidcombe, Australia, 2141; manuscript received 22 April 1987, accepted for publication 18 November 1987. KEY WORDS: Cladonia angustata Nyl., Cladonia bimberiensis A. W. Archer, Cladonia floerkeana (Fr.) Florke, Cladonia macilenta Hoffm., Cladonia murrayi W. Martin, Cladonia pleurota (Florke) Schaerer, Cladonia subdigitata Nyl., Cladonia weymouthi F. Wilson ex A. W. Archer, lichens, Cocciferae, Ochroleucae, chemotaxonomy, distribution, Australian lichens. INTRODUCTION The lichen genus Cladonia Hill ex Browne contrains about 350 species (Ahti, 1982) and of these 53 are thought to occur in Australia. This paper describes the Australian species that form the section Cocczferae (Delise) Dahl, including the group Ochroleucae (Dahl, 1952). The study is based on an examination of relevant type specimens (in- dicated by !), specimens from Australian herbaria, Australian specimens in the British Museum (Natural History) (BM) and the University of Helsinki (H), and the author’s own collections. No previous publication has dealt with this group of lichens in Australia but a recent paper (Stenroos, 1986) has described section Cocczferae from Melanesia, in- cluding Papua New Guinea. The following sixteen taxa of Cladonza sect. Cocciferae have previously been reported from Australia: C. bacillaris Nyl. (Watts, 1903), C. bimbertensis A. W. Archer (Archer, 1985), C. cocczfera (L.) Willd. as Cenomyce coccifera (Brown, 1814), C. corallifera (Kunze) Nyl. (Krempelhuber, 1880), C. cornucopioides (L.) Hoffm. (Leighton, 1867), C. deforms (L.) Hoffm. as Cenomyce deformis (Brown, 1814), C. didyma (Fee) Vainio (Vainio, 1887: 139), C. digitata (Leighton, 1867), C. flabelliformis Vainio (Vainio, 1887: 116), C. floerkeana (Fr.) Florke (Krempelhuber, 1880), C. macilenta Hoffm. (Krempel- huber, 1880), C. murray: W. Martin as Cenomyce firma (Laurer, 1827), C. muscigena Eschw. (Krempelhuber, 1880), C. pleurota (Florke) Schaerer as C. coccifera var. pleurota (Vainio, 1887: 170), C. subdigitata Nyl. (Vainio, 1887: 181) and C. weymouth F. Wilson ex A. W. Archer (Archer, 1985). Several of these reports are not valid and are probably based on misidentifications of other species; eight species in Section Cocciferae are here reported to occur in Australia. A description of each species is given together with its chemistry and distribution. SYSTEMATICS AND DISCUSSION Cladonia sect. Cocciferae (Delise) Dahl Rev. Bryol. Lichenol. 21: 121 (1952) Cenomyce [rankless] Cocciferae Delise in Duby, Bot. gall. 2: 632 (1830). Podetia scyphose or escyphose, simple or slightly branched, often sorediate. Apothecia red, containing rhodocladonic acid, or pale brown. The major phenolic PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 206 THE LICHEN GENUS CLADONIA compounds are usnic, iso-usnic, thamnolic, barbatic, didymic, and squamatic acids; the yellow pigment skyrin and the triterpene zeorin may also be present. KEY to the species of Cladonia sect. Cocciferae in Australia tee Podetiatescy phose years ic See hee here eeu So ares eheoaa eee cls a a 2 Lee Podetias scyphOse (Ms) teh hee tenn Hake lL veut saves beh Ones ne a rr 6 DANS sri Geacid presenti s(t sauce sae sage oe SEO ee tek ce a 3 2 4 Wisnicacid absent: Sys Ni aM oo ce ako We 0 aaah nee, Gl 4 3.) )Rodetia with variablelsonedia: didymic acidi presents). 44))- 5-44 ee C. angustata 3. Podetia with dense farinose soredia; didymic acid absent.......... C. bimbertensis 4. Podetia corticate, or partly corticate and granular sorediate ......... C. floerkeana 4 Podetia parthyiconticate andiwtanrinose sorediate 4 rnans 0-2 ee 5 5. Podetia corticate only at base and just below apothecia; apothecia common; usually Jessttlvara? 25 naaraaatall Ge rear uy oh 7 eh Ee a ae Oh aaa flee C. macilenta 5. Podetia partly corticate, the cortex often covering the lower third of the podetia; apotiecia rare, usuallymoreitham 3 Omraaytall Pssst ee eee eae C. weymouthi 6. Podetia Pd-, thamnolic acid absent; apothecia rare.................. C. pleurota 6. Podetia Pd+ yellow, thamnolic acid present; apotheclacommon.............. ii 7. Usnic acid absent; basal squamules conspicuous, podetia green, growing on soil oda ae aaa ile ccings Sea R Oi a anes MA eMOe ARE NAAN aaa ne Mate ee MAM MEARE CONS oo oc C. murrayt 7. Usnic acid present; basal squamules inconspicuous, podetia yellow, growing on WOO py ayssd ses Sever eal eta a coh Ga ae Re nc ey iti Sere hala aa ne C. subdigitata Cladonia angustata Nyl., Ann. Sci. Nat. Bot. sér. 4, 11: 236 (1859). Cladonia cornucoprordes var. angustata Nyl., Mem. Soc. Scr. Nat. Cherbourg 5: 96 (1858 (1857’) ), nom. nud. Type: Iles Sandwich (Hawaii), J. Remy 1851-1855; lectotype: H-NYL 37978!; isolecto- type: PC. Description: Basal squamules evanescent or persistent, 1 x 1.5mm, lacinate or crenate, esorediate, yellowish-green above, white below. Podetia growing from the basal squamules, 5-10mm tall, 0.5-1.0mm diam., escyphose, simple or rarely branching, apices obtuse or sub-acute, the basal area corticate, the remainder ecorticate and soredi- ate, soredia farinose to granular, cortex sub-continuous or areolate. Apothecia red, terminal. Chemistry: K+ yellow or K-, KC + yellow, P+ yellow or P-. Usnic and didymic acids + thamnolic + barbatic acids. The well-defined basal corticate area on the podetia and the presence of usnic and didymic acids together, distinguish this species from other Australian taxa in the group Cocciferae. Mature apothecia were not seen in Australian specimens. Distribution: Cladonia angustata is an uncommon but widely distributed species in Western Australia, Queensland and Tasmania where it grows on dead wood or sandy peat. It also occurs in New Zealand, Japan and Hawaii. Representative specimens: Western Australia: Busselton, Layman Rd, near Wonnerup House, 31.xii.1981, N. Sammy (PERTH 820560); Porongerups National Park, near summit of Nancy Peak, 7.v11.1968, N. Sammy (PERTH 840899); Jarrahdale, 50km SE of Perth, 9.vi.1973, N. Sammy (PERTH 840934). Queensland: Wybara, 9km NW of Wallanagara, H. Streimann 9829 (CBG). Tasmania: Mawbanna Plain, 15km SE of Stanley, G. Kantvilas 1.iv.1985 (H). PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A.W. ARCHER ZO Cladonia bimberiensis A. W. Archer, Muellerta 6: 93 (1985). Type. Mt Bimberi, 49km SW of Canberra, A.C/T., H. Streimann 9743; holotype: CBG!; isotype: H!, US. Description: Basal squamules persistent, 0.5-1.0mm long, 0.3-0.5mm wide, esorediate, yellow-green above, white below, margins crenate. Podetia growing from the basal squa- mules, 10-30mm tall, 0.7-2.0mm diam., pale yellow, more or less cylindrical, simple and escyphose, or with shallow, deformed scyphi with marginal proliferations; podetia rough corticate at the base and then becoming ecorticate and densely farinose sorediate, with the interior of the scyphi farinose sorediate; esquamulose or occasionally with squamules on the lower part of the podetia. Apothecia not seen; pycnidia subconical, brown, 0.1-0.2mm diam., 0.3-0.4mm long, terminal or marginal on the scyphi; conidia not seen. Chemistry. K-, KC + yellow, P-. Usnic, barbatic and 4-0-demethy] barbatic acids. C. bimberiensis is distinguished from C\ macilenta and C. corniculata, by the yellow colour and the K- and P- reactions, and from C. angustata by the branched podetia and the absence of didymic acid. Distribution: Cladonia bimberiensis is an uncommon alpine to subalpine species found in Victoria, the Australian Capital Territory and ‘Tasmania where it grows on dead wood. It also occurs in the South Island of New Zealand. Representative specimens: Australian Capital Territory: SE of Bimberi Park, Bimberi Range, J. A. Elix 6640 (ANUC, MEL 1047742); ibid. J. A. Elix 6639 (NSW); Mt Frank- lin, A. Archer 1900 (ANUC, H, NSW). Victoria: track to Mt Stirling summit, A. Archer 2058 (NSW). ‘Tasmania: North East Great Lake, G. Bratt 564 (BM). Cladonia floerkeana (Fr.) Florke, Clad. Comment. 99 (1828). ((Floerkiana’). Cenomyce floerkeana Fr., Lich. Suec. Exsic. 82 (1824). Type: not designated. Description: Basal squamules small, inconspicuous, 0.5-1 x 1-2mm slightly lobed. Podetia growing from the basal squamules, simple or sparingly branched near the apices, escyphose, sterile podetia subulate, 5-20(rarely to 25)mm tall, 0.5-l(rarely to 2)mm diam., the major part of the podetia and the area below apothecia corticate, the cortex scabrose to sub-verrucose, the remainder ecorticate and minutely squamulose or granular sorediate, or the podetia completely corticate. Apothecia common, red, convex, terminal, 1-2mm diam. Chemistry: K+ yellow or K-, KC-, P+ yellow or P-. Barbatic and didymic acids + thamnolic acid. Cladonia floerkeana is a widely distributed and, when fertile, conspicuous species in Eastern Australia, distinguished from Cladonia macilenta by the predominantly corticate podetia and the absence of farinose soredia. In contrast to specimens from the Northern Hemisphere the Australian specimens often contain thamnolic acid. Distribution: Cladonia floerkeana is a cosmopolitan species growing on dead or burnt wood or on soil. It occurs in eastern Queensland, New South Wales, Australian Capital Terri- tory, Victoria, South Australia and Tasmania, and also on Norfolk Island. Representative specimens: South Australia: Mt Lofty Range, 5.x.1971, D. Whibley 3678 (AD 97647213). Queensland: 1km S of Herberton, J. Elix 16614 (ANUC). New South Wales: 25km E of Braidwood, 15.x1.1970, E. Dahl (CANB 227864); near Yeomans Bay, ca 30km N of Sydney, 30.11.1985, A. Archer 1741 (H, NSW). Australian Capital Territory: Jervis Bay, H. Streimann 3574 (GBG, H). Victoria: near Beaconsfield, G. Bratt 69/611 (HO 53117). Tasmania: Churchill Spur, Florentine River, G. Bratt 68/237 (HO 53877). Norfolk Island: J. & T. Gilbert, s.n. (HO 53167). PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 208 THE LICHEN GENUS CLADONIA Cladonia macilenta Hoffm. Deutschl. Fl. 2: 126 (1796). Type: not designated. Cladonia bacillaris Ny1., Not. Sallsk. Fauna Fl. Fenn. Forh. 8: 179 (1866). Type: not designated. Description. Basal squamules small, inconspicuous, 0.5 x Imm, margins + soredia. Podetia growing from the basal squamules, simple or rarely branching near the apices, escyphose, pale green, 10-20(rarely to 30)mm tall, 0.5-1.5mm diam., the major part of the podetia ecorticate and farinose sorediate apart from short, smooth corticate areas at the base and below the apothecia; sterile podetia blunt or subulate; rarely squamulose on the basal corticate area. Apothecia red, convex, terminal, 0.5-2mm diam. Chemistry: K+ yellow or K-, KC-, P+ yellow or P-. Barbatic, + didymic + thamnolic acids or rarely squamatic and consquamatic acids. Cladonia macilenta is distinguished from Cladonia weymouth by the short, rarely branched podetia and the small corticate area at the base of the podetia. It is distin- guished from Cladonia floerkeana by the conspicuously ecorticate and farinose sorediate podetia. The exact status of Cladonia bacillaris as a separate species is unclear (Ahti, 1980). Christensen (1987) has recently suggested that Cladonza bacillaris is merely a tham- nolic acid deficient form of Cladonia macilenta. However, in Australia specimens lacking thamnolic acid are uncommon and squamatic acid has only been found in one specimen from Victoria. Distribution: Cladonia macilenta is a cosmopolitan species growing on dead wood or on soil. It occurs in south western Western Australia, eastern Queensland, New South Wales, Australian Capital Territory, and Victoria. Representative specimens. With thamnolic acid: Western Australia: Porongorups National Park, 15.x.1980, N. Sammy (PERTH 810103). Queensland: Tinnaroo Falls Rd, 12km NE of Atherton, J. Elix 16584 (ANUC). New South Wales: Clyde Mtn, 8.xi1.1974, J. Elix (ANUC, MEL 1017163); Bolivia Hill, near Deepwater, 23.1.1980, H. Streimann 9953 (CBG 8002514, H). Victoria: near Moondarra, G. Bratt 76/509 (HO 52909). Lacking thamnolic acid: New South Wales: South Tinderry Peak, 22.11.1970, E. Dahl (CANB 227672); Mt Banks, 9.111.1985, A. Archer 1736 (NSW); near Frederica Falls, Lawson, 27.vi.1987, A. Archer 2115 (NSW). Australian Capital Territory: near Honeysuckle Creek Tracking Station, 2.iv.1983, A. Archer 1483 (NSW). With squamatic acid: Victoria: by the side of Buller Creek, Mirimbula, 5.x1.1986, A. Archer 2049 (H, NSW). Cladonia murrayi W. Martin, Trans Roy. Soc. N.Z., Bot. 2: 40 (1962). Type: Secretary Island, Doubtful Sound, West Otago, New Zealand, Feb. 1959, J. Murray; holotype: CHR 257075!; isotype: BM!. Cenomyce firma Laurer, Linnaea 2: 44 (1827); Cladonia firma (Laurer) Krempelhs, Verh. zool.-bot. Ges. Wien, 18: 309 (1868), nom. illeg.; Cladonia firma (Laurer) Vainio, Acta Soc. Fauna Flora fenn. 4: 215 (1887), nom. inval., non Cladonia firma (Nyl.) Nyl., Bot. Ztg. 47: 352 (1861). Type: New Holland, F. Sieber, sin. loc. [? Tasmania]; holotype: G!. Description: Basal squamules persistent, conspicuous, 1-2mm wide, 3-6mm long, incised, esorediate, the lower surface becoming yellow to yellow-brown in the older parts. Podetia growing from the squamules, 1-2.5cm tall, 1-3mm diam., green or greyish- green, scyphose or with deformed scyphi, with short (2-4mm) proliferations bearing apothecia, corticate, verrucose to scaly, becoming ecorticate and squamulose, esoredi- ate. Apothecia conspicuous, red, 1-3(rarely to5)mm diam. Chemistry: K+ yellow, KC-, P+ yellow. Thamnolic acid and skyrin (rhodophyscin). PROC. LINN. SOC. N.S.W., 110 (2), (1987) 198 A. W. ARCHER 209 Cladonia murrayi is a terricolous, corticate, green, scyphose species with conspicuous red apothecia. It is distinguished from Cladonza subdigitata by the green scyphi and the absence of usnic acid. The exact origin of the type specimen of Cenomyce firma is unclear. The Austrian collector Franz Sieber collected in New South Wales (Audas, 1950: 21) but did not visit Tasmania (Kantvilas, 1983). Recent collections of Cladonia murrayi have been made only in Tasmania and New Zealand and no specimens are known from New South Wales. Herbarium specimens labelled Cladonia murray: from Mt Wilhelm, New Guinea (R. Hnatiuk and E. Dahl, 26.viii.1970) lacked thamnolic acid and skyrin and are identified as Cladonia pseudodigitata Gyeln. (CANB 227807) and Cladonia yunnana (Vainio) des Abb. (CANB 227808). Distribution: Cladonia murray: occurs in Tasmania, growing on moist soil in semi-shaded positions, with Cladia aggregata (Sw.) Nyl. and Siphula decumbens Ny]. It also occurs in New Zealand. Representative specimens: Tasmania: Ben Lomond, May 1887, Dr Bamford (NSW); near summit of Mt Murchison, Dec 1893, Fitzgerald (NSW); Lake Dobson, Mt Field National Park, G. Kantvilas 613/81 (BM); Great Dome, Dennison Range, G. Kantvilas 778/81 (BM); W side of Lake Dove, 30.x1.1983, A. Archer 1562A (H, NSW); 8km from Lake Dobson, G. Bratt 70/303 (HO 56284). Cladonia pleurota (Florke) Schaerer, Enum. lich. eur.: 186 (1850). Capitularia pleurota Florke, Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 2: 218 (1808); Cladonza coccifera var. pleurota (Florke) Schaerer, Lich. helv. spic.: 25 (1823). Type: not designated. Cladonia cornucopioides var. grandis Krempelh., Verh. zool.-bot. Ges. Wien 30: 332 (1881). Type: Mt Ellery, Gippsland, Victoria, 1870, C. Walter; holo: M!. Description: Basal squamules persistent or evanescent, inconspicuous, 1-2mm wide, 2- 4mm long, irregularly crenate to lobate. Podetia growing from the basal squamules, simple, scyphose, scyphi 1-4cm tall, 4-10mm diam, with marginal proliferations with apothecia, or in the form of scyphi; the interior of the scyphi ecorticate with granular soredia or tiny flattened corticate scales; the base of the podetia corticate, the cortex con- tinuous or areolate, becoming verrucose and scaly to subsquamulose, finally ecorticate and granular sorediate in the upper parts. Apothecia red, convex, 1-3mm diam, solitary or clustered on the margins of the scyphi, or pedicellate on the margins of the scyphi, rarely seen in Australian specimens. Chemistry: K-, KC + yellow, P-. Usnic and iso-usnic acids and zeorin. Cladonia pleurota is a common scyphose species and was one of the first Cladonia species collected in Australia by Robert Brown (Crombie, 1880). The presence of iso- usnic acid and the partly corticate scyphi distinguish C. pleurota from all other Australian species of Cladonia whilst the absence of minutely farinose soredia differentiates it from C. deformis. Specimens from Tasmania are often 2-4cm tall. Distribution: Cladonia pleurota is a widely distributed species found on soil in all Australian states except South Australia. It also occurs on Macquarie Island and in New Zealand, Europe, North America and Japan. Representative specimens: Western Australia: Porongorups National Park, 18.x.1980, N. Sammy (PERTH 810019). New South Wales: South Tinderry Peak, 22.111.1970, E. Dahl (CANB 227853); 2km NE of Mt Kosciusko, 14.11.1979, H. Strezmann 7631 (CBG 7907333, H, US). Australian Capital Territory: Black Mountain, 30.vu.1975, B. Hain 40 (CBG 8004587). Victoria: Mt Buffalo, 14.x1.1979, A. Archer 830 (H). Tasmania: Mons Tabulans [Mt Wellington] R. Brown 530 (BM); Arthurs Pass, G. Bratt 72/1810 PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 210 THE LICHEN GENUS CLADONIA (HO 53142). Macquarie Island: 1 mile N of Bauer Bay, 28.1.1964, R. Filson 5812 (MEL 20278). Cladonia subdigitata Nyl., C. r. hebd. Séanc. Acad. Sci., Paris 83: 88 (1876). Type: Campbell Is, New Zealand, 1874, Filhol; lectotype: H-NYL 37858! (T. Ahti, in litt.); isolectotype: MEL!; syntypes: BM!, PC, TUR-V 14155, 14157, nv. Cladonia corallifera (Kunze) Nyl. subsp. subdigitata Vainio, Acta Soc. Fauna Flora fenn. 4: 180 (1887). Type: not designated. Cladonia deformis (L.) Hoffm. var. tasmanica Krempelh., Verh. zool.-bot. Ges. Wien 30: 332 (1881). Type: Tasmania, J. v. B. Gulliver [T. & B. Gulliver]; holotype: M!; isotype: MEL!. Description: Basal squamules pale yellow, persistent, inconspicuous, 1-3 x 1-3mm, subpalmate, margins crenate, esorediate. Podetia growing from the basal squamules, yellow, simple, scyphose, widening gradually from the base, 1.5-3.0(rarely to 5.0)cm tall, 1.5-4mm diam., corticate at the base, becoming scaly or rough or minutely squamulose, corticate below apothecia, scyphi 3-5(rarely to 8)mm diam., interior corticate, closed, or open in older specimens, with marginal scyphi or apothecia. Apothecia red, conspicu- ous, common, convex, 1-4mm diam., marginal on the scyphi or on corticate prolifera- tions from the margins of the scyphi. Chemistry. K+ yellow, KC + yellow, P+ yellow. Usnic and thamnolic acids + skyrin. Cladonia subdigitata is distinguished from all other Australian scyphose Cladonia by the yellow scyphi with conspicuous red apothecia and the presence of thamnolic acid. This taxon is often misidentified as Cladonza coccifera (L.) Willd. on herbaria labels. Distribution: Cladonia subdigitata is a conspicuous, scyphose species growing on dead wood. It occurs in Tasmania and rarely in Victoria, and has been reported from the Grampians, Victoria (Vainio, 1887) and from Papua New Guinea (Mattick, 1942) but this last report was not confirmed (Stenroos, 1986). It also occurs on Macquarie Island and in New Zealand. Representative specimens. Victoria: Lake Tyers, Feb. 1888, F. Wilson (NSW); Mt Macedon, R. Filson 12041 (MEL 39956). Tasmania: Mt Dromedary, June 1895, L. Rodway (NSW); Federation Peak, Jan. 1949, J. Béchervaise (MEL 7106); Mt Mueller, G. Bratt 73/121 (HO 44688); Mt Hartz, G. Bratt 3083 (HO 44721); Lake Fenton, G. Bratt 461 (HO 56247). Macquarie Island: 1 mile N of Bauer Bay, 28.1.1964, R. Filson 5832 (MEL 20277). Cladonia weymouthiu F. Wilson ex A. W. Archer, Muelleria 6: 94 (1985). Type: Huon River, Tasmania, 5 Feb. 1892, W. A. Weymouth; holotype: MEL 6760!; isotype: NSW!. Cladonia cornucopioides L. f. arrosa F. Wilson, Pap. Proc. Roy. Soc. Tasm. 1892-3: 151 (1893). Type: Brown's Track, Mt Wellington, Tasmania, F. Wilson, no date; holotype: NSW L4388!. Description: Basal squamules inconspicuous, persistent, 0.5-lmm wide, 1-2mm long, incised, esorediate, margins crenate. Podetia arising from the basal squamules, 1.5- 5.0cm tall, 1-4mm diam., green to greenish-grey, subcylindrical or tapering to the apices, simple or branching somewhat near the tips, the branching forming deformed scyphi, lacking well-defined scyphi, axils closed; podetia corticate at the base and for one third of the length of the podetia, and below the apothecia, the remainder of the podetia ecorticate and densely farinose sorediate; podetia esquamulose or with squamules on the lower part. Apothecia rare, terminal, red, convex, 1-3mm diam. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. W. ARCHER 211 Chemistry: K- or K+ yellow, KC-, P- or P+ yellow. Thamnolic, barbatic and didymic acids; thamnolic and/or didymic acids may be absent. Cladonia weymouth resembles Cladonia macilenta but is distinguished from that species by the occasional deformed scyphi and the tall, branched, partly corticate podetia. It is distingushed from Cladonza corniculata by the absence of fumarprotocetraric acid. Distribution: Cladonia weymouthi is an uncommon species growing on dead or decayed wood. In Australia it is known only from Tasmania, at altitudes from 250m to 1800m. It also occurs in New Zealand, Papua New Guinea and the Solomon Islands. Representative specimens. Tasmania: Price’s Rivulet, Huon, Feb. 1902, W. A. Weymouth (NSW); near Hastings Cave, 27.x1.1982, A. Archer 1417D (H, MEL 1045447); 15km W of Maydena, 7.x1i.1983, A. Archer 1545A (CBG, NSW); Pencil Pine Creek, 29.xi.1983, A. Archer 1566A (MEL 1045448); the Hermit, 8km SE of Strathgordon, 19.1.1984, G. Kantvilas 59/84 (NSW); Mt Wellington, J. Townrow 34 (HO 54026). EXCLUDED AND DOUBTFUL SPECIES Cladonia bacillaris Ny]. The status of C. bacillaris as a separate species has been discussed by Ahti (1980), Ahti and Stenroos (1986) and Christensen (1987); following Christensen, C. bacillaris is here included in C. macilenta. Cladonia coccifera (L.) Willd. em. Asah. Fl. berol. Prod.: 361 (1787). Lichen cocciferus L., Sp. pl.- 1151 (1753). C. coccifera usually contains usnic and barbatic acids. Although reported to occur in Autralia (Krempelhuber, 1880; Vainio, 1887: 154) no specimens corresponding to C. coccifera have been found. Australian specimens in herbaria labelled ‘C. cocczfera’ are often fertile C. subdigitata but specimens of C. murrayi, C. pleurota and C. merochlorophaea Asah. have also been identified as C. coccifera. Cladonta corallifera (Kunze) Nyl. Flora, Jena 57: 70 (1874). Cenomyce corallifera Kunze, printed herbarium label with description (?1827). C. corallifera, a scyphose species containing usnic, thamnolic and didymic acids, is primarily a lowland Amazonian species from South America (Ahti and Stenroos, 1986). The reported occurrence in Australia is probably based on a misidentification of C. subdigitata, which lacks didymic acid. Cladonia cornucopiordes (L.) Hoffm. Deutsch. Fl. 2: 128 (1796). Lichen cornucoprordes L., Sp. pl. 1151 (1753). C. cornucopiordes is a later synonym of C. coccifera (Vainio, 1887: 149). C. cornucopioides var. grandis Krempelh. (Krempelhuber, 1881) from Victoria was found to be C. pleurota (Acher, 1986b). C. cornucoprordes f. arrosa (Wilson, 1893) from Mt Wellington, Tasmania, NSW 14388, contained only thamnolic acid and is Cladonia weymouthii, vide supra. Cladonia deformis (L.) Hoffm. Deutsch. Fl. 2: 120 (1796). Lichen deformis L., Sp. pl.: 1152 (1753). Cladonia deformis is morphologically similar to C. pleurota and both taxa contain usnic and iso-usnic acids and zeorin. Australian specimens identified as C. deforms, in- cluding the early collection by R. Brown from Tasmania, lack the minutely farinose soredia characteristic of this taxon and these specimens are now referred to C. pleurota. No Australian material resembling northern hemisphere specimens of C. deformis has PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 212 THE LICHEN GENUS CLADONIA been found among the specimens examined. C. deformis var. tasmanica Krempelh. was referred to C. subdigitata (Archer, 1986b). Cladonia didyma (Fée) Vainio. Acta Soc. Fauna Flora fenn. 4: 137 (1887). Scyphophorus didymus Fée, Essai Crypt.: CI (1825). C. didyma, containing thamnolic, barbatic and didymic acids, is a widely distributed pan-tropical to pan-temperate species; the total distribution was recently reported to include Australia (Stenroos, 1986). This record was based on the inclusion of C. didyma in a tentative Key to Australian Cladonia (Archer, 1986a). A re-examination of Australian specimens determined as C. dzdyma has shown that they lack the ecorticate squamulose podetia characteristic of C. didyma so they are now included in C. floerkeana. Cladonia digitata (L.) Hoffm. Deutsch. Fl. 2: 124 (1796). Lichen digitatus L., Sp. pl.: 1152 (1753). C. digitata is a scyphose species, with conspicuous basal squamules and containing thamnolic acid, found in Europe, North America and Japan. No material correspond- ing to C. digitata has been found amongst the collections examined in Australian herbaria. The records of C. digitata from Australia are probably based on misidentifica- tions of C. subdigitata, which contains usnic acid. The detailed description of ‘C. digitata’ provided by Wilson (1893) based on material from Mt Wellington, Tasmania could equally refer to C. subdigitata. A specimen with conspicuous basal squamules and labelled C. digitata from Sandringham, Victoria (NSW) contained atranorin and fumar- protocetraric acid and has been identified as C. praetermissa A. W. Archer. A second scyphose specimen, labelled ‘Cladonia digitata’ from Waterfall, Tasmania (NSW) contains usnic and thamnolic acids and is identified as Cladonia subdigitata. Cladonia flabelliformis Vainio. Acta Soc. Fauna Flora fenn. 4: 113 (1887). C. flabelliformis is a later name for C. polydactyla (Florke) Sprengel (Ahti, 1978). Itisa sorediate, scyphose species containing thamnolic acid, which is widespread in western Europe (Tonsberg and Ahti, 1980). The Australian record is probably based on a misidentification of C. subdigitata as no specimens from Australia have been found corresponding to C. polydactyla. Cladonia muscigena Eschw. In Martius, Fl. Bras. 1 (1): 262 (1833). The type material of C. muscigena was collected in Brazil but the exact status of the species is uncertain (Stenroos, 1986). Vainio (1887) considered C. muscigena to be a variety of C. didyma and it is possible that the first and only report of C. muscigena from Australia was based on a misidentification of C. floerkeana. It is possible that the reports of C. muscigena occurring in New Caledonia are also based on misidentifications of C. floerkeana (Stenroos, 1986). ACKNOWLEDGEMENTS The author is grateful to Professor T. Ahti, Helsinki, for helpful discussion and to the National Herbarium of New South Wales for arranging the loan of specimens. Acknowledgement is made to the Director, Division of Analytical Laboratories, for permission to publish this paper. References AuTI, T., 1978. — Nomenclatural and taxonomic remarks on European species of Cladonta. Ann. bot. fenn. 15: 7-14. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 A. W. ARCHER 213 —, 1980. — Nomenclatural notes on Cladonta species. Lichenologist 12: 125-133. ——., 1982. — Evolutionary trends in Cladoniiform Lichens. /. Hattori Bot. Lab. 52: 331-341. , and STENROOS, S., 1986. — A revision of Cladonia sect. Cocciferrae in the Venuzuelan Andes. Ann. bot. fenn. 23: 229-238. ARCHER, A. W., 1985. — Two new lichens: Cladonza bimberiensis and Cladonia weymouthit. Muelleria 6: 93-95. ——, 1986a. — A tentative key to the lichen genus Cladonza in Australia. 10 pp. Sydney (mimeographed). — , 1986b. — Nomenclatural notes on some Australian Cladonia species. Lichenologist 18: 241-246. AuDAS, J. W., 1950. — The Australian Bushland. Melbourne: Robertson and Mullens Ltd. Brown, R., 1814. — A list of plants natives both of Terra Australia and of Europe. Appendix III, in FLINDERS, M., A voyage to Terra Australis: 592-594. London: G. and W. Nicol. CHRISTENSEN, S., 1987. — Morphological and chemical variation in the Cladonia macilenta/bacillaris aggregate in Denmark. Lichenologist 19: 61-69. CROMBIE, J. M., 1880. — Enumeration of Australian lichens in Herb. Robert Brown (Brit. Mus.), with descriptions of new species. J. Linn. Soc. Lond. (Bot.) 17: 390-401. DAHL, E., 1952. — On the use of chemistry in lichen systematics. Rev. bryol. lichenol. 21: 119-134. KANTVILAS, G., 1983. — A brief history of lichenology in Tasmania. Pap. Proc. Roy. Soc. Tasm. 117: 41-51. KREMPELHUBER, A., 1880., — Lichenes australini e Baronis de Mueller collectionibus. Jn MUELLER, F.,, Fragmenta phytographiae Australiae. Supplementum ad volumen undecimen.: 70-74. Melbourne: Government Printer: ——, 1881. — Ein neuer Beitrag zur Flechtenflora Australiens. Verh. zool.-bot. Ges. Wien 30: 329-342. LAURER, F., 1827. — Sieber’sche Lichenen. Linnaea 2: 38-46. LEIGHTON, W. A., 1867. — Notulae Lichenologicae No. XII. On the Cladonie: in the Hookerian Herbarium at Kew. Ann. Mag. nat. Hist. 19: 99-124. MartTICK, F., 1942. — Die Flechten von Neu-Guinea. 1. Allgemeines. Die Gattung Cladonia. In DIELS, L., Beitrage zur Flora von Papuasien X XVI. Bot. Jb. 72: 151-158. MULLER ARGOVIENSIS, J., 1887. — Revisio Lichenum australiensium Krempelhuberi. Flora, Jena 70: 113-118. STENROOS, S., 1986. — The family Cladonzaceae in Melanesia. 2. Cladonia section Cocciferae. Ann. bot. fenn. 23: 239-250. TONSBERG, T., and AHTI, T., 1980. — Cladonia umbricola, a new lichen species from north west Europe and western North America. Norw. J. Bot. 27: 307-309. VAINIO, E. A., 1887. — Monographia Cladoniarum Universalis. I. Acta Soc. Faun. Fl. Fenn. 4: 1-509. Watts, W. W., 1903. — Notes and Exhibits; twenty-seven lichens from determinations by Dr Bouly de Lesdain of Dunkerque. Proc. Linn. Soc. N.S.W. 28: 498-499. WILSON, F. R. M., 1893. — Tasmanian Lichens Part 1. Pap. Proc. Roy. Soc. Tasm. 1892: 133-178. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 Notes and Discussion A dietary study of Sz/lago analis and its variation in three Australian locations D. T. BREWER AND K. WARBURTON Zoology Dept, University of Queensland, St. Lucia, Australia 4067 The golden-lined whiting, Szl/ago analis (Whitley) is a macrobenthic carnivore widespread around the coast of Australia north of 30°S (McKay, 1985). Two previous studies of the diet of S. analis gave differing results. Lennanton (1969) reported that S. analis, feeding sub-littorally in a mangrove environment in Shark Bay (Western Aus- tralia), showed a strong preference for polychaete worms. Gunn and Milward (1985) found that S. analis caught from the beaches and estuaries around Townsville, fed predominantly on the bivalve mollusc Mesodesma eltane. The present study of S. analis collected from Deception Bay, SE Queensland (27°10’S. 153°00’E) reveals a further variation in the diet of this species. 120 S. analis were collected on six occasions between December and February (summer) of 1983/4, and 200 fish were collected on two occasions in June and July (winter) of 1984. The feeding habitat consisted of an intertidal seagrass flat with a zone of mangroves above mean tide level. Preliminary gut content analyses indicated that S. analis was a consistent nocturnal feeder in this habitat, and therefore collections were carried out at night. Fish were put on ice immediately on capture and had their stomachs excised and preserved in 10% formalin within one hour. Fish ranged in size from 119mm to 322mm (F.L.). Stomach contents were examined using a stereo dissecting microscope where necessary. Most prey animals were identified to genus and many to species. Both numerical and weighted points methods were used to quantify the contribution of prey organisms to the diet of S. analis. The numerical method as described by Hynes (1950) gives an indication of the numbers of each prey type taken by the sampled population. In the case of the weighted points method, the procedure followed was a modification of that described by Hynes (1950). The volume of each food type was estimated by eye and expressed as a percentage of the total stomach contents volume. Items making up less than 5% of the total contents were recorded but were not allocated a points score. The points for each food type were then multiplied by a stomach fullness index assigned using the criteria of Ball (1961) to give weighted points values. Table 1 lists the contribution of prey items to the diet of S. analis at Deception Bay from summer and winter samples. A wide variety of crustaceans was taken; however three crab species (Macropthalmus setosus, M. punctulatus and Ilyograpsis paludicola) predominated. A Wilcoxon test (Siegel, 1956) indicated that there was no significant difference between summer and winter diets. However, Glauconome virens siphon tips, which formed a major part of S. analis’ winter diet, were not seen in summer samples. | Plant material was only seen in small amounts and was probably ingested incidentally with prey items. Gunn and Milward (1985) described the pharangeal dentition of S. analis. They concluded that the broad molariform pharangeal teeth in this species aided in the masti- cation and transport to the oesophagus of both hard and soft bodied prey types. In PROC. LINN. SOC. N.S.W,, 110 (2), (1987) 1988 216 Prey Category CRUSTACEA (a) Natant Decapoda Ocypodidae Grapsidae Portunidae Mictyridae Xanthidae (b) Reptant Decapoda Penaeidae Palaemonidae Callianassidae Alpheidae (c) Isopoda Sphaeromidae (d) Amphipoda Talitridae ANNELIDA Capitellidae Eunicidae Nephtyidae Oweniidae MOLLUSCA (a) Gastropoda Onchidaceae Naticidae Rissoidae (b) Bivalvia Lucinidae Tellinidae Glauconomidae PISCES Gobliidae PLANTS DIETARY STUDY OF SILLAGO ANALIS TABLE 1 The percentage contribution of prey to the diet of Sillago analis at Deception Bay Macropthalmus setosus M. punctulatus Uca sp. TIlyograpsis paludicola Sesarma spp. Portunus pelagicus Thalamita stimpsoni Mictyrus longicarpus Heteropenape sp. Macrobrachium sp. Callianassa australiensis Alpheus edwardsit Onchidium sp. Polinices sordidus Stenothyra sp. Macomona deltoidalis Glaucomone virens UNIDENTIFIED MATERIAL Numerical Method Summer Winter n= 120 n = 200 85.2 48.6 14.6 Teal 21.4 11.6 0.3 0.5 32.5 12.4 3.4 0.6 1.6 0.5 0.4 0.5 0.5 3.8 4.6 fee 0.3 0.1 0.3 4.5 5.4 1.9 1.7 2.6 6.8 3.5) 2.4 1.0 DU 122 0.6 0.7 fel 0.5 1.9 36.2 32 1.3 1.1 1.1 0.3 0.8 0.4 0.6 32.3 0.6 1.8 0.6 1.8 3.8 7.9 125) 1.8 Weighted Points Method Summer Winter n= 120 n = 800 87.3 67.0 21.5 13.4 19.2 14.5 2, 1.3 27.1 11.9 3.6 1.3 1.8 0.7 0.7 1.0 5 32 2.3 1.0 0.2 0.5 0.8 7.4 14.3 3.1 0.6 1.3 5.0 4.6 hel 0.8 3.3 3.4 0.2 0.3 0.4 0.2 2.6 20.8 DoD 5.5 0.1 0.5 0.1 0.4 0.3 0.3 13.9 1.0 4.2 1.0 4.2 2.1 1.4 1.4 1.8) contrast, S. sthama, which occurred sympatrically with S. analzs in the Townsville study, displayed a more delicate pharangeal dentition and was found to eat mainly soft bodied food types (Gunn and Milward, 1985). S. analis therefore appears capable of eating a wide variety of benthic animals. This may be a possible explanation for the variation in the diet of S. analis observed in three different Australian locations. BALL, J. N., 1961. — On the food of the brown trout of Llyn Tegid. Proc. Zool. Soc. Lond. 137: 599-622. References GUNN, J. S., and MILWARD, N. E., 1985. — The food, feeding habits and feeding structures of the whiting species Sillago sthama (Forsskal) and Sillago analis Whitley from Townsville, North Queensland, Aus- tralia. J. Fish. Biol. 26: 411-427. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 D. T. BREWER AND K. WARBURTON 217 Hynes, H. B. N., 1950. — The food of freshwater sticklebacks (Gasterosteus aculeatus and Pygosteus pungitius) with a review of methods used in studies of the food of fishes. J. Anim. Ecol. 19: 36-58. LENNANTON, R. C. J., 1969. — Whiting fishery — Shark Bay. Fishing Industry News Service 2(1): 4-11. W. Aust. Dept Fish. Wildl. McKay, R. J, 1985. — A revision of the fishes of the family Sillaginidae. Mem. Qld Mus. 22(1): 1-73. SIEGEL, S., 1956. — Nonparametric Statistics for the Behavioral Sciences. New York: McGraw Hill. Communicated by J. R. Merrick; manuscript received 5 August 1986, accepted for publication 22 October 1986. PROC. LINN. SOC. N.S.W., 110 (2), (1987) 1988 141 159 175 187 193 205 215 P. GLASBY, P. M. SELKIRK, D. ADAMSON, A. J. DOWNING and D. R. SELKIRK Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western Blue Mountains |. YASSINI and A. J. WRIGHT Distribution and Ecology of Recent Ostracodes (Crustacea) from Port Hacking, New South Wales A T. HOTCHKISS and K. IMAHORI A new Species of Nitella (Characeae) belonging to the Pluricellulate Species Group in Australia A. T. HOTCHKISS and K. IMAHORI Additional Observations on Nitella verticillata (Characeae) from a new Locality in New South Wales R. V. SOUTHCOTT A new Australian larval Callidosomatine Mite (Acarina: Erythraeidae) parasitic on Flies, with Notes on Subfamily and Tribe Classification A. W. ARCHER The Lichen Genus Cladonia Section Cocciferae in Australia Notes and Discussion D. T. BREWER and K. WARBURTON A Dietary Study of Sillago analis and its Variation in three Australian Localities ANNEXURE to nos 1 & 2. THE LINNEAN SOCIETY OF NEW SOUTH WALES. Record of the ANNUAL GENERAL MEETING 1986, Reports and Balance Sheets PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES VOLUME 110 Issued 12th July, 1988 ee ee a CONTENTS: NUMBER 1 1 A. S. STEFFE and B. C. PEASE Diurnal Survey of Ichthyoplankton Abundance, Distribution and Seasonality in Botany Bay, New South Wales : 11 K. G. CAMPBELL and K. M. MOORE Two new Species of Glycaspis (Homoptera: Spondyliaspididae) from potentially endangered Eucalyptus ‘Species and one from E. stricta 19 K.M. MOORE Associations of some Glycaspis Species (Homoptera: Spondyliaspididae) with their Eucalyptus Species Hosts 25 K.M. MOORE A new Species of Glycaspis (Homoptera: Spondyliaspididae) and some new Host Records 27 A. J. WRIGHT First Report of Late Devonian Trilobites from eastern Australia 31. D. K. MCALPINE Studies in Upside-down Flies (Diptera: Neurochaetidae). Part |. Systematics and Phylogeny 59 D. K. MCALPINE Studies in Upside-down Flies (Diptera: Neurochaetidae). Part Il. Biology, Adaptations, and specific Mating Mechanisms 83 F. W. E. ROWE and E. L. ALBERTSON A new Genus and four new Species in the Family Echinasteridae (Echinodermata: Asteroidea) ERRATUM NUMBER 2 101. S. G. M. CARR and D. J. CARR The Sir William Macleay Memorial Lecture 1987. The Elastic-sided Gumleaf, or: the Rubber Cuticle and other Studies of the Corymbosae Contents continued inside a ee TM Printed by Southwood Press Pty Limited, 80-92 Chapel Street, Marrickville 2204 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 110 NUMBER 3 NUMBER 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 1987-88 President: P,. M. MARTIN Vice-president: HELENE A. MARTIN, G. R. PHIPPS, C. N. SMITHERS, A. J. T. WRIGHT Honorary Treasurer? A. RITCHIE Secretary) BARBARA J. STODDARD Council: A. E. J. ANDREWS, M. ARCHER, JUDITH H. K. EASTMAN, R. A. FACER, L. W. G. FILEWOOD, MARILYN D. FOX, L. A. S. JOHNSON; EINE Ary INUAIRSIMING] AR si MIVAURGISTING ) NEB RSRG Cee ee MVERSCOUGEL I> Go PERGIVAL. GRO PHIPES) Aw RIG EM iGemiNe SMITHERS, T. G. VALLANCE, KAREN L. WILSON, A. J. T. WRIGHT Honorary Editor; T. G. VALLANCE — Department of Geology & Geophysics, University of Sydney, Australia, 2006. Linnean Macleay Fellow: JULIA M. HUSH Linnean Macleay Lecturer in Microbiology: KY. CHO Auditors: W. SINCLAIR & CO. The postal address of the Society is P.O. Box 457, Milson’s Point 2061, N.S.W., Australia. Telephone (02) 929 0253. © Linnean Society of New South Wales Cover motif: ‘Fossils of Talbragar’, a vignette designed and drawn for this volume by Alan E. J. Andrews. PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES | Marine Biotogical Go APR 26 ggg Woods Hole, Mass VOLUME 110 NUMBER 3 7 ant et Wise Fluxes of Inorganic Nitrogen between Sediments and Water in a Coral Reef Lagoon R. JOHNSTONE*, K. KOOP and A. W. D. LARKUM JOHNSTONE, R., Koop, K., & LARKuM, A. W. D. Fluxes of inorganic nitrogen between sediments and water in a coral reef lagoon. Proc. Linn. Soc. N.S.W. 110 (3), (1988) 1989: 219-227. Lagoon sediments of coral reefs are potentially important sites of nutrient regeneration. Profiles of ammonium within sediments at One Tree Island, southern Great Barrier Reef, showed strong gradients with values as high as 50ug atoms NH,*-N.1! at 15cm depth decreasing to around 8yg atoms.!" just below the surface. Gradients of NO3;/NO, also existed but concentrations were much lower than for ammonium, ranging from undetectable levels to approx. 8ug atoms.l'. Overlying waters characteristically have low to undetectable levels of dissolved ammonium or nitrate/nitrite. Nutrient profiles were examined in the top 0-10cm of two sediment types and despite strong concentration gradients, no efflux of NH,* ions or NO3 /NO, ions were observed from the sediment into the water column. In experiments where the water under domes was artificially enriched with ammonium chloride to maximum sediment concentrations (50ug atoms NH,*-N.I') rapid uptake of NH, *-N by sedi- ments was shown to occur. Microbial processes such as nitrification, denitrification, and ammonium utilization by psammolithic algae in the uppermost layers of sediment may be responsible for the observed phenomena. R. Johnstone and A. W. D. Larkum, School of Biological Sciences, University of Sydney, Sydney, Australia 2006, and K. Koop, Zoology Department, Stockholm University, S-106 91, Stockholm, Sweden; (R. Johnstone, present address, Zoology Department, Stockholm University, S-106 91, Stockholm, Sweden); manuscript received 17 March 1987, accepted for publication 23 March 1988. INTRODUCTION Since the pioneering work of Odum and Odum (1955), Wiebe e¢ al. (1975), and Webb et al. (1975), different aspects of nitrogen dynamics on coral reef ecosystems have been investigated (e.g. Wiebe, 1976; Smith and Jokiel, 1975; Webb and Wiebe, 1975; Capone et al., 1975; Hatcher and Hatcher, 1981; and Koop and Larkum, 1987). Apart from the work of Entsch e¢ al. (1983) and Williams et al. (1985), however, there are few published studies on nitrogen cycling in coral reef sediments, although many such studies exist for other environments (see e.g. Raine and Patching, 1980; Hopkinson and Wetzel, 1982). Nitrogen fixation is widely believed to be responsible for supplying large amounts of nitrogen to coral reef ecosystems (Webb et al., 1975; Wilkinson et al., 1984; and others) although Koop and Larkum (1988) have shown that only 10-15% of nitrogen require- ments of primary producers at One Tree Island, southern Great Barrier Reef, can be met by direct nitrogen fixation. There is still a substantial deficit which must be met from other sources such as remineralization by microorganisms. This study assesses the role of sediments as a source of recycled nitrogen for the adjacent systems by measuring levels of ammonium, nitrate and nitrite within sedi- ments as well as fluxes of these ions between sediments and the overlying water column in One Tree Is. lagoon, southern Great Barrier Reef. ee re a nNN Bi SPREE cL Oe ere aE 3 Bet al Aisa ied Crete DE Te Su ee Bh ae a * Formerly, Linnean Macleay Fellow. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 220 FLUXES OF INORGANIC NITROGEN MATERIALS AND METHODS Sites The whole of this work was conducted in the main lagoon on One Tree Island Reef, Great Barrier Reef, Australia (lat. 20°30°S, long. 152°06’E), (Fig. 1) during summer, 1984. Nutrient fluxes were measured in the two sediment types representing the range in sediment environments found in One Tree [s. lagoon. These were very fine sand and coarse sand with >40% of particles <63um diameter and >40% of particles >1mm diameter respectively. TOWNSVILLE QUEENSLAND ° 144 Fig. 1. One Tree Island reef, southern Great Barrier Reef showing (B, right) experimental sites in two sediment types. © Coarse sand; * Very fine sand. Mean water depth was 4.5m at the site with very fine sand and 1.5m at the site with coarse sand. All sediments in One Tree lagoon contain approx. 97% calcium carbonate, derived from the fringing and patch reefs. Pore Water Sampling and Analysis Four replicate cores 5cm diameter and 20cm deep, were taken randomly in areas of each sediment type and subsamples of 10cm? were taken at depths of 0.5, 2.5, 5.0, 10.0 and 15.0cm from each core. The pore water from each subsample was extracted by centrifugation at 3,000 x g for 10 minutes and the concentration of ammonium, nitrate and nitrite ions determined according to Strickland and Parsons (1972). All analyses were conducted within 2 hours of core collection. Nitrate and nitrite analysis was conducted using a Jechnicon auto-analyser, model AA1. Rates of efflux of NH,* from the sediments were calculated using the equations of Rutgers van der Loeff et al. , (1984): PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 R. JOHNSTONE, K. KOOP AND A. W. D. LARKUM 221 J) = @D), leks (Equation 1) where J, = xateof flux; g = sediment porosity, D, = effective diffusion coefficient, dC/dx = concentration gradient across the sediment/water interface. Measurement of Sediment Fluxes Measurement of fluxes of ions from the sediment were carried out using six poly- carbonate domes placed randomly over the sediment at each site. The domes had a volume of 36 litres with a basal area of 640cm? and were anchored to a steel skirt pushed 5cm into the sediment. The skirt also served to prevent lateral movement of water under the edge of the domes. In each dome, water above the sediment was sampled every 2 hours for 18 hours, using a 20ml syringe and needle pushed through a self-sealing sampling port on the side of the dome. An equal amount of seawater was allowed to enter the dome during sampling to prevent water being drawn from the sediment itself. impeller——__ sampling port sealing gasket steel skirt Fig. 2. Dome enclosures used for flux measurements and enrichment experiments showing hand driven impeller, sampling ports and anchoring base. The water inside each dome was stirred with a hand-driven impeller mounted in its centre (Fig. 2) for 5 minutes every hour, and just prior to sampling. Care was taken to ensure that the stirring rate was low enough to leave the sediment undisturbed, but strong enough to ensure thorough mixing of the dome water. Samples were immediately refrigerated for transport to the laboratory and analysis for ammonium, nitrate and nitrite ions was carried out within 5 hours of collection as for pore-water samples. Enrichment Experiments In experiments using four replicate domes placed randomly over the sediment at each site, the water under the domes was enriched with ammonium chloride to produce a concentration of dissolved ammonium ions approximately equal to the maximum concentration found in the underlying sediment. Four control domes were also set up at PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 Dodge FLUXES OF INORGANIC NITROGEN each site with no enrichment. Water in all domes and their immediate surrounds was sampled every 2 hours for 17 hours and the samples analysed for ammonium concen- tration, oxygen concentration, pH, and temperature. Also, as a control for uptake of ammonium by organisms in the water column, twelve 500ml bottles, 6 light-proof and 6 transparent, were filled with seawater taken at each site and enriched with ammonium chloride to the same concentration as the en- riched domes. Controls were incubated zn sztu for the duration of the enrichment experi- ment and sampled at the start, midway, and at the end of the experiment. The samples were analysed for dissolved ammonium as before. Temperature and dissolved oxygen were measured zn situ with a submersible com- bination temperature/oxygen probe attached to a YSI meter (model number 57; Yellow Springs Instruments, Ohio, U.S.A.). Sample pH was determined immediately after collection with an Activon 105 portable pH meter. The expected flux rates of ammonium ions from the enriched dome water into the underlying sediment were calculated with the Stokes-Einstein equation for diffusion (Crank et al., 1981): M/2.(aDt)'/?.exp(-r?/4Dt) (Equation 2) = concentration of solute, quantity of solute emitted at t=O from unit area of the plane of origin, = the diffusion coefficient, = (ahene, = the distance of the point of measurement from the plane of source. where ll This equation describes the decrease of a solute with time due to simple diffusion. By considering the underlying sediment as an infinite sink the equation would give the maximum rate of diffusion possible and enable us to speculate on the possibility of active uptake by sediment or water column organisms. RESULTS Pore-Water Ammonium, Nitrate/Nitrite, and Sediment-Water Column Fluxes Both sediment types showed the same concentration of free ammonium at 15cm depth (38ug atoms NH,*-N.1") but above this depth the concentration gradients were considerably different (Fig. 3). In the very fine sand site the concentration had a pronounced peak at 5cm depth (50ug atoms NH, *-N.1"') with the lowest concentration in the top 1cm of sediment (18g atoms NH,t-N.171). By comparison the coarse sand site showed a peak ammonium concentration at 15cm depth (38ug atoms NH,*-N.1"), with a gradual decrease toward the top 1cm of sediment (8ug atoms NH,+-N.1I"), (Fig. 3). Nitrate/nitrite concentration gradients for both sediment types showed a small peak just below the sediment surface; 6.75 + 8.8ug atoms NO;/NO,-N.1! (n=4) at 5cm in the coarse sand and 7.0 + 7.8ug atoms NO3/NO,-N.1'! (n=4) at 2.5cm in the very fine sand (Fig. 4). In both sediment types, NO3;/NO, concentrations decreased with in- creasing sediment depth and concentrations were usually barely above detection limits. Despite the existence of reasonably strong concentration gradients of NH,* and the occurrence of measurable amounts of NO; /NO, in the sediments at One Tree Is., no flux of dissolved NH,* or NO; /NOz into the overlying water column was observed from either type of sediment studied. Also no dissolved ammonium was detected in the surrounding seawater at the site with coarse sand, but water at the site with very fine sand consistently had an NH,+ concentration of 1.9 + 0.1pmols.1!. NO3/NO, ions were undetectable in the surrounding water over both study sites. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 R. JOHNSTONE, K. KOOP AND A. W. D. LARKUM 223 60 50 40 (pg atN.I") 36 10 0 5 10 15 20 Depth (cm) Fig. 3. Concentration gradients of ammonium in pore-water of two sediment types at One Tree Is. Vertical bars are 95% confidence limits (n=4). e@ecoce Very fine sand; — — —--- Coarse sand. Using the concentration gradients from the top 5cm of each sediment type the expected rates of NH,* efflux from the sediment surface were calculated with the Stokes-Einstein equation. This was corrected for temperature variation and used a diffusion coefficient of 19.8 x 10°cm?.s! (Li and Gregory, 1974). The resulting flux rates were 2.55ug atoms N.m~.h? for coarse sand, and 1.85y4g atoms N.m~?.h! for the very fine sand. 10 oO 5 10 15 Depth (cm) Fig. 4. Concentration gradients of nitrate/nitrite in pore-water of two sediment types at One Tree Is. Vertical bars are 95% confidence limits (n=4). O-———-—O Very fine sand; ¢ © Coarse sand. PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 Doers FLUXES OF INORGANIC NITROGEN Enrichment Experiments Experiments in sites with both the very fine sand and coarse sand showed approxi- mately exponential loss of NH,* over the experimental period (Fig. 5A,B). Above the very fine sand the rate of loss showed a short but distinct plateau centred on the time of total dark, approximately 7.5 hours after the start of the experiment. The plateau lasted approximately 4 hours and was followed by a slower rate of decrease in NH,* ions than previously observed. Above the coarse sand there was no plateau, and the decrease in ammonium ion concentration over time was approximately exponential (Fig. 5B). 50 40 + 30 (y mol ") 20 2 4 6 8 10 12 «#414 16 18 2 4 6 8 10 12 #14 6 18 Time th) Time th) Fig. 5. Concentrations of ammonium in enriched dome water over very fine sand (A), and coarse sand (B) at One Tree Is. Note the plateau between 6 and 10 hours at the very fine sand site. Total dark occurred at 7 hours for both experiments. Vertical bars are 95% confidence limits (n=4). Using the Stokes-Einstein equation for diffusion, the calculated concentrations of ammonium ions in the water under the domes after 17 hours would be 10uM NH,* and 12uM NH,* for very fine and coarse sands respectively (this assumes diffusion at the sediment surface into an infinite sink for NH,*). The concentrations observed after 17 hours were 10 + 2uM for very fine sand and 5 + 2.5uM for coarse sand. These values are equal to or less than those calculated, indicating that the ammonium ions were being taken up at near maximal rates. At the end of the experimental period, there was no significant difference between individual domes in the two types of sediments, or between treatments, for mean values of pH, temperature or dissolved oxygen in the water enclosed by the domes, and there was no difference between entrapped dome water and unenclosed water for these parameters (Table 1). Also, there was no decrease in the concentration of ammonium ions in the control bottles of enriched water incubated over the 17 hour period indicating that there was no detectable uptake of ammonium ions by water column organisms. DISCUSSION Physical variations, such as grain size, have received little attention in relation to nutrient levels in sediments of coral reefs (e.g. Entsch et al., 1983). We have shown here (Fig. 3) that the ammonium concentration in the first 15cm of sediment is higher in the fine compared to the coarse sediment. Although many factors may be involved in deter- mining these gradients, it is likely that differences in the metabolic activity of microbial populations of each sediment type varies the nutrient levels. This conclusion is PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 R. JOHNSTONE, K. KOOP AND A. W. D. LARKUM TABLE 1 225 Mean values and standard deviations of dissolved oxygen, pH, and temperature in enriched dome water, control dome water, and surrounding seawater over the two study sites for enrichment experimen:s January, 1985 O, pH ‘Temperature (ppm) (°C) Very Fine Sand. Enriched a8) es Ilo72) 8.03 + 0.09 28.25 + 0.82 Control DROZ ple23 8.06 + 0.10 28.25 +0.86 Seawater 6.17 + 0.73 8.08 + 0.18 28.57 + 0.90 Coarse Sand. Enriched FBT 23 MSF 8.01 + 0.18 27.88 + 1.16 Control 6.87 + 2.00 7.99 + 0.18 27.95 + 1.24 Seawater 6.20 + 1.51 7.96 + 0.14 28.00 + 1.45 consistent with work elsewhere that shows grain size and porosity influence the nutrient levels of sediments, and are correlated with changes in sediment microbial and in- vertebrate populations (Dale, 1974; Jansson, 1967). The concentration gradients from NO;/NO,° observed here are similar to those found in coral reef sediments elsewhere (Corredor and Morell, 1985) and the concen- trations found at One Tree Is. fall in the same range as found in sediments on the north- ern Great Barrier Reef (Boon, 1986). We suggest that the peak in NO;/NO. concentrations close to the sediment surface is due to bacterial nitrification which has been shown in other types of marine sediment to occur in the oxygenated zone (e.g. Henriksen, 1980; Henriksen et al., 1981). Ammonium needed for this process would be supplied from the deeper anoxic zones where ammonification occurs and the concen- tration gradients observed here are more than adequate for this process. The absence of a NO; /NO, flux from the sediments into the water column is consistent with work on fluxes by Harrison (1983), who found that they were mostly negative (i.e. from the water column into the sediment). It is likely that the NO3/NOz,° pool, which is small com- pared with the ammonium pool is either utilized as a nitrogen source, or used in denitrification by the microbial population in the top layers of sediment. For dissolved ammonium, concentration gradients and pools were up to an order of magnitude higher than for NO;/NO,. Despite the relatively steep concentration gradients of dissolved ammonium ions, and although an efflux of NH,* ions has been observed elsewhere for coral reef sediments (Williams et al., 1985), no efflux of ammonium ions from the sediment into the water column was observed in this study. Using the NH,* concentration gradients in the One Tree reef sediments, and assuming a passive system, there should be a theoretical flux of NH,4* ions out of the sediment of 2.55umols NH,*+-N.m?.h7, for coarse sand; and 1.85umols NH,+-N.m™!.h", for very fine sand. The fact that no such flux was observed indicates that an active uptake process must be occurring in the top layers of sediment and this is substantiated by the fact that when the water under the domes was enriched with ammonium ions, its concentration decreased over time despite an opposing concentration gradient in the underlying sediments. Applying the Stokes-Einstein equation for diffusion (Crank et al., 1981) of ammonium ions out of the domes and into the sediment, it is possible to show that ammonium was leaving the dome at near maximum rates (assuming a diffusive process). Ammonium ions are not used by organisms in the water column as shown by the bottle experiments, thus it is concluded that the ammonium is rapidly taken up and used, or modified, by organisms in or on the top layers of sediment; despite the fact that these organisms are also receiving a flux of ammonium ions from deeper sediments. PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 226 FLUXES OF INORGANIC NITROGEN Use of free ammonium ions by marine microalgae is well known (e.g. Syrett, 1953; Goldman and Glibert, 1982) and microscopic inspection of water from the enriched domes and the top Icm of sediment showed a suite of microalgae to be present in the sediment, but only rarely present in water samples. Thus microalgae may at least in part be responsible for the uptake of ammonium from the sediment but as indicated above other processes such as nitrification and denitrification may also change ammonium concentrations. Because of the size of the ammonium pool in One Tree reef sediments, however, it is unlikely that it could all be processed via nitrification and subsequent denitrification. If this were the case, then the corresponding denitrification rate would be approximately 2.4umols N,O.m~.h"!. This is almost ten times the rate observed by Seitzinger and D’Elia (1985) for coral reef sediments (0.3umols NyO.m*?.h"), and com- parable to rates found in marine sediments with a much higher organic input (Nishio et al., 1983). The level of nitrification and denitrification in One Tree reef sediments is presently being studied. Apart from microbial composition, other biological factors likely to affect sediment nutrient levels are bioturbation and grazing which occur in surface layers. Both types of sediment discussed here are disturbed by burrowing shrimps and deposit feeding organ- isms such as holothuria. Such organisms significantly affect bacteria in sediments (Juniper, 1981). At One Tree reef, most turbating organisms were observed in the top 15cm of sediment where the concentration gradients between the two sediment types are most different. In addition, the coarse sand is often exposed to currents which can totally rework the top 5 to 10cm of sediment (Frith, 1985). Although such currents were not observed during the period of this study, they occur fairly frequently. Combined with the activi- ties of turbating organisms such currents probably affect the incorporation of organic matter into, and general stability of the sediment. In summary these results show that the sediments at One Tree reef do not release free dissolved ammonium, nitrate or nitrite ions for use by primary producers or other organisms in the water column or hard substrate communities, but appear to contain microorganisms capable of using and modifying this nitrogen. Presumably nitrogen ultimately leaves the sediments in forms other than dissolved ions possibly via grazers of micro-organisms, but the processes involved are still to be identified. ACKNOWLEDGEMENTS We thank Philip Smith and Penny Butcher, managers of One Tree Island field station, as well as Rod Smith for their support and assistance in the field. The study was supported by the award of a Linnean Macleay Fellowship and a Joyce Vickery research grant from the Linnean Society of N.S.W. to R. Johnstone, and an Australian Marine Science and Technologies grant to A. W. D. Larkum. References Boon, P. L, 1986. — Uptake and release of nitrogen compounds in coral reef and seagrass, Thalassia hemprichit (Ehrenb.) Aschers., bed sediments at Lizard Island, Queensland. Aust. J. Mar. Freshw. Res. 37: 11-19. CAPONE, D. G., TayLor, D. L., and TayLor, B. F., 1975. — Nitrogen fixation associated with macroalgae in a coral reef community in the Bahamas. Mar. Biol. 40: 29-32. CORREDOR, J. E., and MORELL, J., 1985. — Inorganic nitrogen in coral reef sediments. Mar. Chem. 16: 379-384. CRANK, J., MCFARLANE, N. R., Newsy, J. C., PATERSON, G. D., and PEDLEY, J. B., 1981. — Diffusion Processes in Environmental Systems. London: Macmillan Press Ltd. DALE, N. G., 1974. — Bacteria in intertidal sediments: Factors related to their distribution. Limnol. 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Marine Ecology (in press). Li, Y.-H., and GREGORY, S., 1974. — Diffusion of ions in seawater and in deep-sea sediments. Geochim. Cosmochim. Acta 38: 703-714. NIsHIO, T., KOIKE, I., and HATTORI, A., 1983. — Estimates of denitrification and nitrification in coastal and estuarine sediments. Appl. Environ. Microbiol. 45: 444-450. OpuM, H. T., and OpuM, E. P., 1955. — Trophic structure and productivity of a windward coral reef com- munity on Eniwetok Atoll. Ecol. Monogr. 25: 291-320. RAINE, R. C. T., and PATCHING, J. W., 1980. — Aspects of carbon and nitrogen cycling in a shallow marine environment. J. Exp. Mar. Biol. Ecol. 47: 127-139. RUTGERS VAN DER LOEFF, M. M., ANDERSON, L. G., HALL, P. O., IVERFELDT, A., JOSEFSON, A. B., SUNDBY, B., and WESTERLUND, S. F. G., 1984. — The asphyxiation technique: An approach to distinguish between molecular diffusion and biologically mediated transport at the sediment-water interface. Limnol. Oceanogr. 29: 675-686. SEITZINGER, S. P., and D’ELIA, P., 1985. — Preliminary studies of denitrification on a coral reef. The Ecology of Coral Reefs. REAKA, M. L. (ed.). NOAA Undersea Res. Ser. SMITH, S. V., and JOKIEL, P. L., 1975. — Water composition and biogeochemical gradients in the Canton Atoll lagoon: 2). Budgets of phosphorous, nitrogen, carbon dioxide and particulate material. Mar. Scz. Comm. 1: 162-207. STRICKLAND, J. D. H., and PARSONS, T. R., 1972. — A practical handbook of seawater analysis (2nd ed.). Fish. Res. Bd. Can. Bull. 167. SYRETT, P. J., 1953. — The assimilation of ammonia by nitrogen starved cells of Chlorella vulgaris. Ann. Bot. (Lond.) 65: 1-19. WEBB, K. L., and WIEBE, W. J., 1975. — Nitrification on a coral reef. Can. J. Microbiol. 21: 1427-1431. ——, DuPauL, W. D., WIEBE, W. J., SOTTILE, W., and JOHANNES, R. E., 1975. — Enewetok Atoll: Aspects of the nitrogen cycle on a coral reef. Limnol. Oceanogr. 20: 198-210. WIEBE, W. J., 1976. — Nitrogen cycle on a coral reef. Micronesia 12: 23-26. ——,, JOHANNES, R. E., and WEBB, K. L., 1975. — Nitrogen fixation in a coral reef community. Sczence 188: 257-259. WILKINSON, C. R., WILLIAMS, D. McCB., SAMMARCO, P. W., Hocc, R. W., and Trott, L. A., 1984. — Rates of nitrogen fixation on coral reefs across the continental shelf of the central Great Barrier Reef. Mar. Biol. 80: 255-262. WILLIAMS, S. L., GILL, I. P., and YARISH, S. H., 1985. — Nitrogen cycling in backreef sediments; St. Croix, USS. Virgin Is. Proc. 5th Int. Coral Reef Congress, Tahiti. 3: 389-394. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 fi ie Day d tag aH; ; bie a sey id oor i apy ie mh | ihn ivf ate i GR} Hy see | ne ales on mit Asset wees te ie aids i ' sabniee ihe by a CAN fen ‘ erry : mane Bi vaio an ru wiry ky (ing) fan ceuite: rae 1. “heed shen) eee bce teh i} tg MOP ie Pia meted ’ at soe, og Alan. e: “ry ial x rey 2 eee) at con ane ea aud 4 i 1K Rath tae BR iA 4 er dcns a ied i 2 Ea i a — hu J te . yi ort> & ij = bef ¥ wy Liar | walled ih oor: ATT, Nida 16-4 £54450 BIR sy ie. L J i iy: Avrd pla ily ty be ohinedh 4 j ' b ae j rhutet rt - i i ¥y Lh rR Ms hac f i b hhess Z : ‘ : Hos, GLARE pee 3 Vara iy ah hs (ae eis . xi 14 bey ‘ y om - a hd LA) ha af : ae ; ’ Ai ; att Hirer? 1a) . ce 5 ‘ d - uit a ‘ 5 " st 7 4 we * i yy) ' a ‘ a 4 Y - i 5 Le 2 bs : P 7 Pe it r zs Sf 4 OO ec) Ie iii, Dt vA ei Estuarine Foraminiferal Communities in Lake Illawarra, N.S.W. I. YASSINI and B. G. JONES YASSINI, I., & JONES, B. G. Estuarine foraminiferal communities in Lake Illawarra, N.S.W. Proc. Linn. Soc. N.S.W. 110 (3), (1988) 1989: 229-266. Lake Illawarra is a shallow brackish coastal lagoon which formed behind a marine sand barrier following the last rise in ocean level. It is connected to the ocean by an active tidal channel. Tidal flushing of the lagoon is slow and it ranges in salinity from 16°/o0 to 40°/oo. The lagoon is rapidly filling with sand and silt, with the sand mainly confined to banks around the lagoon margins. The substrate and turbidity of the lagoon control the distribution of floral communities which, in turn, influence the benthic invertebrates. The distribution and composition of estuarine foraminiferal communities in Lake Illawarra depend on salinity, substrate and the rate of tidal exchange. In semi-enclosed estuarine lagoons the diversity of foraminiferal assemblages shows a marked decrease from the open ocean to the body of the lagoon. The connecting channel between the lagoon and the ocean is subject to diurnal tidal flushing and it contains a mixed assemb- lage of reworked and living intertidal and subtidal species with minor contributions of lagoonal species. The assemblage in the mobile sands of the tidal channel is much lower than on the seagrass-covered banks of the channel. Within the main body of the lagoon, where salinity is slightly lower than in the open ocean and tidal exchange is minor, foraminiferal assemblages are controlled by the substrate and its associated floral con- stituents. The sandy sediments around the margin of the lagoon have a foraminiferal assemblage characterized by a range of textulariinid species whereas the deeper and silty portions of the lagoon are dominated by two species of rotaliinids. Ammonia beccari is an environmentally-tolerant species occurring at almost all the lagoonal to intertidal marine sample sites. Foraminifer population densities in the lagoon range from high to very low or absent depending on the quantity of available oxygen at the sediment-water interface. In New South Wales, lagoonal foraminiferal communities differ significantly from communities developed in more open estuaries where tidal exchange is rapid. In the latter foraminiferal communities are more diverse and depend largely on the sub- strate and water depth, with salinity variations only affecting populations in the upstream portions of the estuary. I. Yassini and B. G. Jones, Department of Geology, University of Wollongong, PO. Box 1144, Wollongong, Australia 2500, manuscript received 17 June 1987, accepted for publication 23 March 1988. INTRODUCTION The distribution of foraminifers in Lake Illawarra, a coastal lagoon located 80km south of Sydney in New South Wales, has been determined from 132 samples collected from the lagoon, its tidal entrance channel and the adjacent marine intertidal Windang Island. The purpose of this study is to describe the faunal composition and distribution patterns of the foraminifers in the lagoon and adjacent upper intertidal environments. Foraminiferal assemblages have been described from a number of bays, estuaries and lagoons along the coast of New South Wales (e.g., Albani, 1968, 1978, 1979, 1981; Albani and Johnson, 1976; Cotter, 1980) and from other Australian coastal environ- ments (e.g., Apthorpe, 1980; Cann and Gostin, 1985; Collins 1958, 1974; Parr, 1932, 1945; Quilty, 1976). In addition, a number of studies have provided information about the physical parameters and seagrass communities in N.S.W. estuarine environments (e.g., Anderson and Story, 1981; Anderson et al., 1981; Eliot et al., 1976; Harris 1976a, b; Jones et al., 1976; Public Works Department, 1985; Roy and Peat, 1975; Yassini and Wright, 1988) which can be related to their foraminiferal assemblages and thus provide a basis for erecting general foraminiferal biotope models for coastal environments. _ PROC. LINN. SOG. N.S.W., 110 (3), (1988) 1989 230 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PHYSICAL CHARACTER OF LAKE ILLAWARRA A recent appraisal of the physical and chemical character of Lake Illawarra has been given in Yassini and Jones (1987). Lake Illawarra is a shallow elongate coastal lagoon situated 80km south of Sydney, N.S.W. It covers an area of about 34km? (Fig. 1) and fills a coastal depression scoured into the Late Permian Shoalhaven Group. PORT KEMBLA f MULLET f/ @ sample site * no forams >50% sand FSH fe @ 4 TALLAWARRA //@. 159 / @ 158 Entrance channel WINDANG ISLAND ¥ 40-52 MACQUARIE RIVULET BGJ/DMM/GD797 B Fig. 1. Sample locations, water depths (in metres) and the distribution of sand-dominated substrates in Lake [llawarra. The lagoon was formed during the last rise in ocean level some 6000-6500 years ago by the formation of a marine sand barrier north and south of Windang Island which im- pounded and flooded the former shallow marine embayment. The lagoon has a maxi- mum depth of about 3.7m and an average depth of 1.9m (Roy and Peat, 1975). At present the lagoon water level is about 25-30cm above the mean ocean level and it is PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 231 separated from the ocean by a tidal channel 2.4km long, approximately 600m wide and up to 3.5m deep. The seaward end of the channel is continuously changing its position and shape, and heavy shoaling of the mouth occasionally causes complete closure of the lagoon. Lake Illawarra has an average annual fresh water input of about 80 x 10°m? (Ellis et al., 1977). The tidal range along the landward margins of the lagoon is about 3-5cm increasing to 10-25cm near the tidal entrance channel. The average tidal exchange between the lagoon and the ocean is about 0.5 x 10°m? giving a residence time of approximately 26-39 weeks for the lake water mass (Ellis et al., 1977). Tidal currents are essentially limited to the entrance channel and currents within Lake Illawarra are mainly wind generated (Clarke and Eliot, 1984). The average monthly lagoon surface water temperature ranges from 9°C in June to 29°C in February. At most times there is no significant difference between the surface and bottom water temperatures in the lagoon. Sandy sediments within Lake Illawarra are confined to water depths shallower than 2m and they are most abundant along the eastern margin of the lake and around the Macquarie Rivulet and Mullet Creek deltas (Fig. 1). The central part of the lagoon has low relief with a depth of 2m to 3.7m and the sediments consist of dark grey silt and clay containing volatile organic matter (15-19%), iron monosulphides and faecal pellets. CHEMICAL CHARACTERISTICS OF LAKE ILLAWARRA The salinity of the lagoon ranges from 16°/oo to 40°/oo depending on the balance between fresh water inflow, evaporation and the rate of tidal exchange. The salinity is within the normal estuarine range and, except during flood periods, low salinities are confined to stream mouths. Throughout the main body of the lagoon there is very little lateral or vertical variation in salinity values (<3°/oo, Ellis et al., 1977) due to wind induced circulation and mixing of the water. The mean pH of the surface layer in the lagoon varies between 7.3-8.3. In late winter, spring and summer, blooms of macro-algae increase the surface pH to almost 10 during the day but it returns to 7.3 at night. However, just below the water-sediment interface the connate water may be slightly acidic. According to Ellis and Kanamori (1977) and the Electricity Commission of New South Wales (1982-1983) the concentration of dissolved oxygen in the surface layer of the lagoon during the day varies between 4-11mg/l. However, large fluctuations in dissolved oxygen concentrations between night and day occur in areas on the eastern side of the lagoon affected by algal blooms. For example, during one bloom in Griffins Bay the dis- solved oxygen content ranged from a daytime maximum of 14mg/] to a night-time mini- mum of 1.5mg/I, with only 0.5mg/I of dissolved oxygen at the water-sediment interface. Salinity stratification and oxygen depletion of the bottom layer often occur after major flooding and periods of heavy rain (Gibbs, 1986). Nocturnal oxygen deficiencies are important in pH-Eh-related reactions in the bottom sediments, especially in the release of phosphorus to the water column and the denitrification of nitrate and nitrite. As an enclosed saline coastal lagoon, with very limited tidal exchange and almost entirely surrounded by residential and industrial development, Lake [lawarra is sub- jected to continuous pollution. The pollutants are of three major categories: suspended solids, excess nutrients and heavy metals. Suspended solids consist mainly of silt and clay which are removed from the catch- ment by erosion, especially river bank collapse, and are washed out of urban centres bringing some 100,000m? of sediments annually into the lagoon. This can rise to 270,000m? in years with major floods (Hean and Nanson, 1985). Flood tidal currents PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 232 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA bring an annual average 110,00m° of marine sand (Public Works Department, N.S.W., 1985) into the lagoon through the entrance channel. Fly-ash particles from Tallawarra Power Station (1952-present) and Port Kembla (1928-present) are also abundant in the lagoon sediments (Jones et al., 1976). Coal-wash and slag deposited as back-fill in the lagoon catchment area have also contributed to the solid pollutants in the lagoon. Sus- pended solids increase turbidity, inhibit photosynthetic processes and increase the rapid infilling of the lagoon. The mean turbidity level of the lake for the period 1984-1985 was ONTU. Rural and urban drainage networks contribute large quantities of nitrogenous and phosphatic leachates to the lagoon and its bottom sediment and these contribute to the excess growth of macro-algae. During June 1985 an algal bloom produced 71,000 tonnes of macro-algae (Yassini, in prep.) and the subsequent decomposition of this mass caused an oxygen deficiency at the sediment-water interface allowing the production of H)S and black iron monosulphides. High localized concentrations of heavy metals in the top 25cm of lagoon sediments have been recorded by Jones et al. (1976), Roy and Peat (1975) and Ellis and Kanamori (1977). The main pollutants are zinc, lead and cadmium which were probably derived from the Dapto Smelting Works (1895-1906) and the Port Kembla industrial complex since 1928. BENTHIC FLORA AND FAUNA IN LAKE ILLAWARRA Both angiosperms and algae are important benthic floral constituents in the lagoon but are confined to water depths shallower than 2m since the high turbidity restricts light penetration to greater depths. On the shallow water sandy and muddy sandbanks along the tidal entrance channel and Windang Peninsula, around Mullet Creek and Macquarie Rivulet deltas and in Griffins and Koona bays four species of aquatic angio- sperm (Ruppia megacarpa Masson, Zostera capricornt Aschers., Halophila decipiens Ostenfeld and Halophila ovalis (R.Br.) Hook. f.) are abundant. They are associated with green, red and brown algae, and several species of epiphytic blue-green and red algae occur on the leaves of Zostera and Ruppia. On rocky outcrops along the northern, western and southern shores of Lake Illawarra brown, green and red algae are the dominant floral constituents. Details of the algal flora are given in Yassini and Jones (1987). The benthic invertebrate fauna in Lake Illawarra displays a low diversity but a high population density which is typical of estuarine systems. Polychaete worms and the molluscs Yéllina (Macoma) deltoidalis (Lamarck), Spisula trigonella (Lamarck) and Hydrobia buccinoides (Hedley) form over 90% of macrobenthic biomass. Other constituents include additional molluscs, amphipods, isopods, cirripedes, ostracods, decapods, reptantes and tubellarids. FORAMINIFERS For each of the 132 samples used in this study about 100cc of sediment was wet- sieved on a 200-mesh screen and the foraminifers were collected from 50cc of the washed residue, using conventional micro-palaeontological techniques. The foraminifers were not stained for live specimens since 2/3 of the samples came from a previous study (Jones et al., 1976). The foraminifer classification (Appendix) used in this study conforms mainly to that of Loeblich and Tappan (1964, 1974, 1984). Additional refer- ences which were very useful for the specific identification of these southeastern Aus- tralian foraminifers include Albani (1974), Barker (1960), Brady (1884), Chapman and Parr (1937), Cushman (1932, 1933, 1942), Hermelin and Scott (1985), Kohl (1985), PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 233 Matoba (1970), McCulloch (1977, 1981), Murray (1975), Poag (1981), Seibold (1975) and Sidebottom (1912, 1913). A total of 123 benthonic and 6 planktonic foraminiferal species have been recog- nized in Lake Illawarra, its tidal entrance channel and the intertidal zone surrounding Windang Island. All the benthonic species are shallow water forms and the vast majority of these species have been recorded from the diverse populations at Windang Island (85 species) and the entrance channel to the lagoon (89 species). Within Lake Illawarra 16 species have been recorded, mainly from the margins of the lagoon; only two species are common from the central part of the lagoon. Thirty nine samples from the lagoon and entrance channel contained no foraminifers. Distribution of all identified species is summarized in ‘Tables 1 to 3 and in the Appendix. The total benthonic foraminiferal population density is indicated in Fig. 2. TABLE 1 Samples containing only Ammonia beccarii and Cribrononion sydneyensis Ammonia beccari only 239, ¥Y23, Y26, Y29, Y34, Y36 Ammonia: Cribrononion ratio >10:1 77, 162, 467, Y24 10:1 to 5:1 340, 376, 386, Y6, Y7 5:1 to 2:1 161, 164, 166, 187, 198, 203, 220, 236, 342, SOS Ss NOL NCE Ne NAO NOE NEI 2:1to1:1 21,87, 142, 158, 159, 169, 181, 202, 234, 262, 265, 384, Y16, Y22, Y25, Y31, Y32, Y39 <1:1 85, 112, 310, 359 The foraminiferal population within the lagoon is dominated by two of the three species of rotaliinids with more restricted distributions for the two species of miliolinids and eleven species of textulariinids. The rotaliinids form the most abundant group in the lagoon and are represented by Ammonia beccarit and Cribrononion sydneyensis (Tables 1 and 2). A comparison of Figs 1 and 3 indicates that the two dominant species of rotaliinids in the lagoon constitute the entire foraminiferal population throughout almost all the deeper parts of the lagoon and its shallow embayments where the substrate is muddy. Crzbrononion sydneyensis is confined to the deeper water areas (Fig. 4) whereas Ammonia beccarii is a very tolerant species occur- ring in almost all foraminiferal assemblages from the lagoon (Table 1), tidal channel and intertidal environments. It dominates at water depths of less than 1m where the sub- strate is sandy but it decreases in relative abundance from the lagoon towards the open ocean. Textulariinids are most abundant around Mullet Creek and Macquarie Rivulet deltas, Griffins Bay and the tidal entrance channel (Fig. 5). Agglutinated foraminifers were mainly observed on sandy substrates but in Griffins Bay they were present in clay- rich sediment. While there is no distinct areal zonation of textulariinid species through the lagoon there is a distinct difference between the lagoon and the entrance channel- marine assemblage. The textulariinid species confined to the lagoon include Protoschista findens, Miliammina fusca, Ammobaculites agglutinans, A. foliaceus, Texturalia porrecta, E-ggerella australis and Trochammina inflata. Tritaxis conica occurs both in the northern half of the lake and in the entrance channel and marine assemblages. Mailiolinids have a very restricted distribution within the lagoon system and are confined essentially to the entrance channel, the shallow zone along Windang Peninsula and isolated occurrences around delta mouths (Fig. 6). The two species, Trzloculina oblonga and Miliolinella subrotunda, were found on sandy substrates with dense to PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA 234 = ls {8s €°GG = OT = sc = i= = = = = = = = 8x = CUE. 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SNuDIUOSWDIVIM saprourMaNg S1)19D4B SaproulMnNg psojnurgs vjyjassnay DIDUIBIDU DUIWANG DIDINGNS DjDSUOJa DUIWLAING DINIDIL]S DUIVZUG DIDID DUIDZuG “ds vuzarjog DISNgOL DUIAIOg pyvajdgopnasd pur21j0g wunjo1ofiag vuranog izatuop DuIIxn0g DjIvGuo0) vur1j0g I] ‘ds 0u270¢ ] ‘ds vuzj0Q putas DID}IUNGO}DIL}8 DUI0CQ DjDIUI) DUILOQ suazog viupjnuvjy DJDIINS DUIBDT DISOINJNIVGNS DUIS] DIVILISIUAIS “JI DUIBDT DJDaUIPWIS DUIDD] DIVINGUA “JI DUISDT DULY IVT “JI DUIGDT DULI}}1IVDLG DUIGDT DIDUILI DUIDD'T DISOINNID DUIS] [] ‘ds vuunssr7 [ ‘ds vurunss.y DIVING VULENSSV vypiponbgns “JI DUUNSSIT DIDLOf{LIGOJDUITADUL DULANSS Lo] DIDUNID) DULUNSSL] DIDUILDI DIDIISV{ DUINSSL] PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 DSi) I. YASSINI AND B. G. JONES 74 “a = € 08! gI = 0z = = 01 02 1€ a6 = = OE! - = Ol 6£ = = = = = = 01 6% = = Ol 6z = = 01 8L = 02 z 09 6 08 06 = = 0s 08! Ol = 02z 8 = = b 001 01 OF 001 = rs = El = = = a = z = € = = = 9 = = 81 cl ~ - - = = = 01 98 Ge € 01 ZI = = € G = = z GGA 90K box GIN pues] Suepulny = b = = ¢ @ = = = = € = = = = = OE! = 01 02 ¢ = = = 09 = 9 I z = z = = 02 = = 01 = = 7 = = 001 € 02 OL 0¢ = ¢ = ¢ = € G @ = = = = ¢ = € ¢ z = = = = 01 = = € = = * == = — = —_ G — — = = = 0% = = 0¢ + = = = = OLI = 8 01 01 = = = 0% = = cI z = = = = = = = = = z = = 91 = = = = = = = = 082 € GE = rl 0€ 6S = 9 b = = ¢ = = = = = 9 = = = = = = = — — — ¢ — — —_— — — 02 = = = = = = = = G — — — _ — — — — cI = = = = = = = = — = = G = =e — — — GZ = = 01 I = = = = ¢ —_— — — —_— — — —_ — = = = G — — — — = € = = = € = Z = = IPA 66A OCGA 6IA 8IA L6G 686 ELE 6S Jauueyp dduReTUq (penunuod) ¢ ATV snynssasgap UOiUon) XUDLAGULL DUIPIALAD J ‘ds wnipiyd) 7 wnpjarvu wnipryd) 5] tuasual wnipryd) 7 unjnssasdap wnipryd)y wndsi19 unipryd) q Snaquasiv mnipryd) J unuaapp wnipryd) sisuadaupds woruouoLsgusy SISUIUDLLI}IPAUL DUIINGLOUD] J suasavyur DUIINALIIP wdpo.g vjjaLogojvqud’) “ds puryrs0Un 7] upog DuLvSOY S1]DAJSND DUIIVDSOY DINTUD DUILDSOY snanjuong saprournunid mng1dsuosur vjjaurjajvd SN]DIPLULIp S1gLOISIpOpjauD’] vanquos0un)g 07]aU2gGL0IS1CT 170]AYIL9q D]JaUIgLOSSIC vsojnisng vyja2mng sisuaurddipyg vuisdog I] ds myja7psqnps) ] ‘ds oyjarnsqnjy SUDINIDULAGD] D]JA]DLQDIL) pyourajnd 0jja7vLqG0] 5) srusofrjajod 0)]3j049)]) wLog “J vjjaqosqnjy) S1SUA]DLISND D)]A]DLGD]£) suvjnsuniponb srgs09sipojnsuy pursasag “JI DuLLasIA/) sisuassng DurLas10/) PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA 238 b = I = t 4 z = € = 4 4 = vjvjfur (vyyvjo10gin J) v170}04090).) a = = = ¢ = _ I = = = = _ DINSLLY 017D}0409 0] =a as = = = = = = Zz = = = = pjojnzojanbygo vurjviuayjng aa a = = <= = Zz I = _ = = = pssaaiun DUIINGLE) = 22 = = = = CG Zz = = = = _ tadj4ajnp vuisponboqgo)s) Or = 7 ag Or = & S = = a = = 4aqna saprours992g0)) Ol = = = = = = = = = = — — “ds nu299190/5) = = — S Or ay 4 £ G = I — = Sap2oy]ng vur499190)5) = = = = a = = I = = = = — “ds punjiids #E oe = = = = = = = = Or = = pivgiara vurppr4igy - ast = = = _ = I — = = = = DjvjnI49gn} DUIpLALIGS = = oa $ = = = = = = = — = syonbanur vurpirg¢ = = = = a = = Zz = = = — = DjD]NIYUaP DUIP)LULIGS 8] = = fee = = = = = = _ = = DJVONLLOI DUIP}IIDY = = = = = = = C Zz I EI = = saprouoiuou DurvUaUp 09 = aS Or Or = $ = C = = = = supprsasig sap19igi20dq 0g = = = = = ae = — = _ = = suadpnfas sapi9ig?) 98 = G O€ OF G Or <4 Or F rea = = wunsoudd? sapingi) 8 bent = * = ae = ss = = = = = ppinj42g 01]0}0Y4 0d — C 0 if 0Z 6 CE 08 OS —_— _ OL _ 41LDII9Q vi UdULET 4 or — — 0 if 0 I — _— 0 I _ _ —_ I _ Sud nv D))]2U0r uON 66A OCGA 6TA BIA L6é G86 €L6 6S Jauueyp souRnUA oGA 90% boA GbX ThA purysy Suepurry (papnpuos) ¢ ATAVL PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 239 PORT KEMBLA f GRIFFINS BAY MULLET 0 L 2km CREEK @® sample site * no forams PURRY-BUARY POINT TALLAWARRA POWER STATION e —~\ iy / Bevins \e'0° Jy Fs e / ISLAND Entrance channel + WINDANG ISLAND MACQUARIE RIVULET Fig. 2. Total number of benthonic foraminifers in 100cc of sediment at each sample site. moderate sea grass coverage in the northern and southern parts of the lagoon respec- tively (thus accounting for their poor correlation with the more uniformly distributed textulariinid species in the sandy lagoon margin sediments — Fig. 7). The entrance channel and Windang Island samples contain a mixed assemblage of intertidal and subtidal benthonic foraminifers and a few planktonic species (Table 3; Fig. 8). This is a typical intertidal population which is similar to many other areas on the N.SW. coast (e.g., Albani, 1968, 1979; Albani and Johnson, 1976). Foraminifers from a constant volume of each sample from the area studied were subjected to R-mode and Q-mode multivariate analysis. Based on similarity levels and grouping of samples (Figs 7-8) the following four distinct assemblages (Fig. 9) were recognized within the studied area. PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 240 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PORT KEMBLA f IY? GRIFFINS ; B f, ) Kin ate i. tel e ° ; 50 2 7 fi I : (ee ea) CG e eas e ® sample site \\ | 50 2 taal, e ® ° %* no forams 0 . 50 a ; ¥ Y) \ e PURRY —-BURRY att ~——-> POINT 7 ir | @ / x et es Oey arenes 0 > (es ( * Tv TALLAWARRA Bs \F ey POWER STATION z Ve oN oO — Be oS* \ 2 ® © / 19 ue e ly * WINDANG Cr e | BEVINS a O)) / ISLAND }'sQ_ 2 e@ iL Tie a : [ 6 . 5 @' Entrance \w e e channel A ,. WINDANG 4 ee AO PRP ISLAND =e a ee a 50 < uy @ ro) = ‘ N S W 2 = e 9° | f S \ | . eu e i Li ae : BAY \ Fig. 3. Distribution of rotaliinids in Lake Illawarra (in percent). Assemblage I This assemblage (27 samples) occurs in the deepest parts of the lake on a silty clay substrate. It is dominated by Ammonia beccari (52.7%) and Cribrononion sydneyensis (43.9%) with subordinate Triloculina oblonga (3.0%). Another six species of textularinids and miliolinids comprise less than 0.4% of the total fauna in this assemblage and mainly occur in samples from close to the lake margin. Assemblage II The 37 samples comprising Assemblage II have even more limited foraminiferal diversity. It is dominated by Ammonia beccaru (83.2%) and Cribrononion sydneyensis (16.4% ). Only two other species occur in this assemblage, namely Ammobaculites foliaceus (0.4%) and Miliolinella subrotunda (0.03%). PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 241 PORT KEMBLA f MULLET 0 L 2km CREEK @ sample site * no forams TALLAWARRA POWER STATION © 5 rays Entrance channel = WINDANG ae Re ISLAND MACQUARIE RIVULET Fig. 4. Distribution of Cribrononion sydneyensis in Lake Ilawarra (in percent). Assemblage III Assemblage III occurs in the four areas around the margins of the lake where nutrient supplies are greatest (that is, adjacent to the tidal entrance channel), Macquarie Rivulet and Mullet Creek deltas and Griffins Bay. In all these areas the sub- strate is predominantly sand and the flora is dominated by Zostera capricorn: Aschers. ‘The 15 samples are dominated by textulariinids (10 species, 49.6%), rotalids (6 species, 44.8%) and Miliolinella subrotunda (5.6%). The main textulariinid species are Ammobacu- lites folraceus (18.2%), Tritaxis conica (10.5%), Trochammina inflata (7.6%) and Eggerella australis (6.3%) and the two dominant rotaliinids are the lagoonal Ammonia beccariu (33.3%) and Cribrononion sydneyensis (10.5%). Assemblage IV The fourth assemblage consists of a mixture of predominantly marine foraminifers (Fig. 8), some of which represent a life assemblage. Approximately half of the species PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 242 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PORT KEMBLA f MULLET ¥ Oar, e) 2kM Creek x bd - a [a = — * 350 SLE IES (a> =~ EWG e | @ sample site DUR IN ( SES | * no forams ® \ \ e \\ pam 0) l) \ Xe 0 2 SS Ly WA PURRY-BUARY —— a ga = POINT * * @ e\e N e e e * e * e a~ = (CoN an nye TALLAWARRA ~ 4 oN O POWER STATION a e Ns \ i © ° “ i) A ee ie * WINDANG ft oO BEVINS p x e ISLAND 0 ~. 5 e i Wee | * by i e © Entrance . e e ® channel ze.) WINDANG & Cm) : : AO°} ISLAND ra F Cer so oo A: 0 < wy e i . B> e : NS W He 0 e 0 ! = le O e Be | * Sydney XN N — Ven nw Fig. 5. Distribution of textulariinids in Lake Illawarra (in percent). probably have been reworked from subtidal environments into Assemblage IV and are only represented by a few specimens. The tidal entrance channel apd Windang Island areas have 55 species in common and of these 35 species probably represent live species based on more than 8 specimens being present at a sample site. The most abundant species constituting an average of 70% of the population at each site in Assemblage IV are: Mean number Mean % abundance of individuals at each location Lamellodiscorbis dimidiatus 215)59) Poff | Rosalina anglica 54.8 5.0 Elphidium crispum 410).7/ 9.0 Rosalina bradyi 36.0 3.9 Elphidium macellum veg 3.6 Rosalina australis 27.6 2.8 Quinqueloculina subpolygona Zeal 4.6 Ammonia beccarit 18.8 Tal Cibicides cygnorum 16.7 4.6 Glabratella australensis 13e1 De PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 243 PORT KEMBLA f GRIFFINS BAY MULLET 0 | 2km CREEK @ sample site * no forams PURRY-BUARY POINT TALLAWARRA POWER STATION HAY WARDS Entrance channel WINDANG Os Re ISLAND MACQUARIE RIVULET Fig. 6. Distribution of miliolinids in Lake Illawarra (in percent). The mixed nature of this tidal assemblage 1s clearly illustrated in Fig. 8 and consists of. lagoonal, intertidal and subtidal species forming a complex thanatocoenose. Minor subdivisions of this assemblage are dependent upon preferential association with par- ticular algal species, especially coralline algae and Zonarza. LAKE ILLAWARRA FORAMINIFERAL COMMUNITIES The assemblages in Lake Illawarra and its tidal entrance channel are very typical of estuarine lagoons with restricted tidal circulation (e.g., Apthorpe, 1980; Cotter, 1980; Michie, 1982; and Murray, 1973). The distribution pattern and species diversity of foraminifers in the Lake Illawarra area can be used as a basis for distinguishing mud- and sand-dominated lagoonal environments from tidal channel and more open estua- rine bay environments such as Broken Bay (Albani, 1978) and Botany Bay (Albani, 1981). The foraminiferal assemblages in Lake Illawarra can be directly related to the floral communities with the greatest diversity of foraminifers occurring in the seagrass beds PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 244 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA -0.2 0 0.2 0.4 0.6 0.8 1.0 Miliammina fusca Trochammina inflata Protoschista findens Ammobaculites foliaceus Tritaxis conica Ammobaculites agglutinans Texturalia porrecta Eggerella australis Sandy lagoon margins Miliolinella subrotunda Triloculina oblonga Ammonia beccarii Deeper part of lagoon Cribrononion sydneyensis Fig. 7. R-mode cluster dendrogram for species occurring at more than two localities within Lake Illawarra. Ammonia beccarit and Cribrononion sydneyensis form a distinct group characteristic of deeper water or silty sedi- ments whereas the textulariinids characterize shallow sandy substrates. where there is relatively low turbidity, a low mud content and an abundant oxygen supply. However, these areas are subject to wider fluctuations in pH and Eh, as well as moderate variations in salinity and temperature, compared with the main body of the lagoon where circulation is less restricted. Foraminiferal faunas on the lagoon margins are dominated by typical euryhaline eurythermal lagoonal species such as Cribrononion sydneyensis, Ammonia beccarit, Miliolinella subrotunda and textulariinids (Fig. 7; Table 2). In the higher energy, wave-influenced margins of the shallow lagoon sandflats the substrate is too mobile for colonization by and preservation of foraminifers and most samples in these regions were barren. In the deeper portion of Lake Illawarra and in sheltered shallower regions mud is the dominant substrate and, together with the relatively high turbidity, prevents the establishment of seagrass beds. In these areas the foraminiferal diversity is very low (only one or two dominant rotaliinid species) but the population densities range up to 350 specimens per 100cc of sample, the highest density in the lagoon system (Fig. 2). A few shallow water areas within the muddy facies have large accumulations of decaying organic matter (e.g., parts of Griffins, Haywards and Koona bays) resulting in an- aerobic bottom conditions which inhibit benthonic foraminifers colonizing these areas. This assemblage therefore shows a great range in foraminiferal abundance which is dependent on a number of environmental factors, not just salinity variation. The tidal entrance channel to the lagoon is characterized by a very diverse total foraminiferal population with a mixed assemblage of both marine and lagoonal species. Few foraminiferal species live in the mobile sandy tidal channel, which contains a partially abraded mixed marine and estuarine fauna, whereas the seagrass banks along the channel margins support a diverse assemblage with strong shallow marine affinities. The only major distinction between this facies and intertidal open marine facies is the greater abundance of reworked lagoonal species in the tidal channel (although they form less than 2% of the foraminifera population in the channel area). PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 Dominantly Mixed tidal channel and shallow marine fauna tidal channel and lagoon I. YASSINI AND B. G. JONES 0.25 0.5 0.75 1.0 [o) Dominantly intertidal to sublittoral marine Miliolinella circularis Dyocibicides biserialis Fissurina fasciata carinata Parrellina imperatrix Gaudryina convexa Angulodiscorbis quadrangularis Planorbulina mediterranensis Triloculina trigonula Cymbaloporetta bradyi Quingueloculina subpolygona Miliolinella baragwanathi Glabratella australensis Quinqueloculina poeyana Guttulina pacifica Fissurina lacunata Buliminoides gracilis Haplophragmoides canariensis Acervulina inhaerens Cibicides cygnorum Quinqueloculina pseudoreticulata Elphidium crispum Nonionella auris Globigerinoides ruber Baggina phillippinensis Elphidium macellum Elphidium depressulum Uviverina bassensis Globorotalia (Turborotalia) infl Brizalina alata Textularia sagittula Textularia candeiana Glabratella tabernacularis Rosalina anglica Triloculina oblonga Globigerina bulloides Elphidium jenseni Rosalina bradyi Lamellodiscorbis dimidiatus Patellinella inconspicua Rosalina australis Anomalina nonionoides Brizalina striatula Elphidium advenum Tritaxis conica Amphicoryna scalaris Ammonia beccarii 245 ata Fig. 8. R-mode cluster dendrogram for species occurring at more than two localities in the tidal channel and around Windang Island. A closely correlated, dominantly marine fauna shows strong similarity to the mixed assemblage of the tidal channel whereas these assemblages show very poor correlation with the lagoonal to tidal channel group. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 246 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PORT KEMBLA f GRIFFINS BAY MULLET 0 I 2k ™M GREEK @® sample site * no forams PURRY-BURRY TALLAWARRA Entrance channel x WINDANG oe Re ISLAND MACQUARIE RIVULET Fig. 9. Foraminiferal assemblages in Lake Illawarra defined by Q-mode cluster analysis. ESTUARINE FORAMINIFERAL COMMUNITIES IN N.S.W. Distribution patterns and species diversity of foraminiferal faunas in the estuarine environments along the coast of New South Wales show two distinct ecological situ- ations. The first is a low diversity and low to moderate population density model, typi- fied by examples of shallow coastal lagoons such as Lake Ilawarra, Narrabeen Lagoon, Tuggerah Lake and Lake Macquarie. The second type exhibits high-diversity and medium-density populations in drowned valleys and estuaries, illustrated by example of Broken Bay, Botany Bay and Jervis Bay. In both models the mixing rate of fresh and saline water masses, and salinity fluctuations, play a determinant role in the faunal dis- tribution. The dynamic setting of the environment and the mobility of the substrate are the main factors controlling faunal distributions within the different salinity zones. Sand-barrier Coastal Lagoon Foraminiferal Community In N.S.W. lagoons such as Lake Illawarra, Narrabeen Lagoon and Tuggerah Lake the fine-grained sediment in the main body of the lagoon supports a dominant assem- PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 247 blage of Ammonia beccarit and Cribrononion sydneyensis together with a few arenaceous species such as Eggerella australis, Textularia porrecta and Ammobaculites agglutinans. Similar assemblages were also observed in Lake Victoria and the Gippsland lake system (Apthorpe, 1980), and in Hardy Inlet, Western Australia (Quilty, 1976). The seagrass beds in the peripheral zone around the margin of these lagoonal environments supports a fauna dominated by arenaceous forms such as Trochammina inflata, Ammobaculites agelutinans and Eggerella australis and a few species of Miliolidae (e.g., Muliolinella subrotunda and Triloculina oblonga). The tidal channels connecting these lagoons to the open ocean contain a mixture of reworked forms from the nearby oceanic intertidal zone and the lagoon proper. In Lake Illawarra, 82 species were encountered in the seagrass beds along the banks of the inlet channel where the energy level is more reduced. In the middle of the channel the mobile sandy substrate contains only a minor abraded and reworked microfauna. Hardy Inlet contained 42 species in the connection channel (Quilty, 1976). In Narrabeen Lagoon 38 species were identified in the inlet channel while the main body of the lagoon contains only 12 species (Yassini, in prep.). Tidal Estuary Foraminiferal Community In the second estuarine ecological type, where tidal exchange of the watermass 1s rapid, the diversity and density of the foraminiferal population gradually decreases from the mouth of the bay towards the higher reaches of the estuary. In Broken Bay, Albani (1978) recorded 181 species of benthonic foraminifers along 40km of estuary. The density of the foraminifer population was low both at the upstream end of the estuary and at the outer entrance of the estuary. Upstream in the non-tidal section of the estuary, where the fresh and saline water masses meet and oligohaline con- ditions prevail, only a few arenaceous foraminifers were found. In the main section of the estuary where there is little change in salinity, the fauna is mostly dominated by Miliolidae (31 species) and Nodosariidae (23 species). Where the estuary has a restricted oceanic connection, foraminifer populations decrease in diversity due to the mobility of the sandy substrate during each tidal cycle. In the Port Hacking estuary, Albani (1968) recorded 119 species of foraminifers along a 10.4km stretch of the estuary. In the weakly-tidal fluvial-dominated section of the estuary only 15 species were found. A maximum diversity of 108 species was found in the main tide-dominated section of the estuary where Miliolidae were represented by 18 species (only 15% of the total fauna). Based on both foraminifer and ostracod distri- butions, the ecological model in Port Hacking is intermediate between a typical open oceanic bay and a lagoon (Yassini and Wright, 1988). Open Ocean Foraminiferal Communities In the upper intertidal zone along the foreshore of Windang Island near the oceanic end of the entrance channel to Lake Illawarra, the foraminiferal density and diversity are typical of an open ocean embayment environment. In the zone 85 species were recorded with Nodosariidae (8 species) and Miliolidae (16 species) forming a prominent component of the foraminiferal assemblage. Foraminiferal assemblages in the open ocean are controlled by the substrate, energy and depth of the environment but a discussion of these facies is beyond the scope of this paper. ACKNOWLEDGEMENTS The authors wish to acknowledge their indebtedness to Mrs Anne Clarke, Secre- tary of the Lake Hlawarra Management Committee, and to the Management Com- mittee for their support and encouragement. Dr A. J. Wright reviewed the manuscript PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA 248 * a Sead e as bee » a bal 2h eto eo Rs rae m Me > LINN PROC I. YASSINI AND B. G. JONES 249 and provided many useft:l suggestions. Dr A. D. Albani checked some of the species identifications. Most of the SEM micrography was kindly undertaken by the electron microscope unit at the University of Sydney. The use of facilities and technical assistance from the Department of Geology, University of Wollongong, are gratefully acknowledged. Grants from the Illawarra Region of Councils (1984-1985 Regional Community Development Programme) and from the University of Wollongong enabled this study to be undertaken. References ALBANI, A. D., 1968. — Recent Foraminiferida from Port Hacking, New South Wales. Contr. Cushman Fdn foramin. Res. 19: 85-119. ——, 1974. — New benthonic Foraminiferida from Australian waters. /. foramin. Res. 4: 33-39. ——,, 1978. — Recent Foraminifera of an estuarine environment in Broken Bay, New South Wales. Aust. //. mar. Freshwat. Res. 29: 355-398. — , 1979. — Recent shallow water Foraminifera from New South Wales. Australian Marine Science Association Handbook 3: 1-57. ——., 1981. — Pleistocene Foraminifera from Botany Bay, New South Wales. Alcheringa 5: 147-160. , and JOHNSON, K. R., 1976. — Resolution of foraminiferal biotopes in Broken Bay, New South Wales. J. geol. Soc. Aust. 21: 435-446. ANDERSON, J. R., and Story, K. J., 1981. — Temperature, salinity, turbidity and tidal data for eight estuaries on the south coast of New South Wales, 1974-1978. Tech. Memo. Commonw. scient. ind. Res. Org. 81/20. , and CAROLANE, R., 1981. — Macrobenthic fauna and sediment data for eight estuaries on the south coast of New South Wales. Tech. Memo. Commonw. scient. ind. Res. Org. 81/22. APTHORPE, M., 1980. — Foraminiferal distribution in the estuarine Gippsland Lakes system, Victoria. Proc. Roy. Soc. Vict. 91: 207-232. BARKER, R. W., 1960. — Taxonomic notes on the species figured by H. B. Brady in his report on the Foraminifera dredged by H.M.S. Challenger during the years 1873-1876. Spec. Publs Soc. econ. Paleont. Miner., Tulsa 9: 1-238. Braby, H. B., 1884. — Report on the Foraminifera dredged by H.M.S. Challenger during the years 1873- 1876. Challenger Expedition Report, Zoology 9: 1-814. CANN, J. H., and GosTIN, V. A., 1985. — Coastal sedimentary facies and foraminiferal biofacies of the St. Kilda Formation at Port Gawler, South Australia. Trans Roy. Soc. S. Aust. 109: 121-142. ? CarRTER, A. N., 1964. — Tertiary Foraminifera from Gippsland, Victoria, and their stratigraphical significance. Mem. geol. Surv. Vict. 23: 1-154. CxiarRKE, D. J., and Exror, I. G., 1984. — Wind driven water circulation in Lake Illawarra, New South Wales: a numerical simulation. University of Wollongong Coastal Research Report 2/84. CHAPMAN, F., and PARR, W. J., 1937. — Foraminifera. Australian Antarctic Expedition 1911-1914 Scientific Report Series G. COLLINS, A. C., 1958. — Foraminifera. Great Barrier Reef Expedition 1928-1929 Report 6: 335-436. ——.,, 1974. — Port Phillip survey 1957-1963, Foraminiferida. Mem. Nat. Mus. Vict. 35: 1-62. CorTtTer, L. K., 1980. — The distribution of Foraminiferida in the Clyde River estuary, Batemans Bay, N.S.W. North Ryde, N.S.W.: Macquarie University, B.A. (Hons) thesis, unpubl. CUSHMAN, J. A., 1932. — The Foraminifera of the tropical Pacific collections of the Albatross, 1899-1900. Bull. U.S. natn. Mus. 161, Pt 1. Fig. 10. 1-3, Trochammina inflata Sample 245, Lake Ilawarra. x54 Spiral, apertural and umbilical views; 4, Ammobaculites foliaceus Sample Y15, Lake Illawarra. x50 Spiral view; 5, Ammobaculites agglutinans Sample Y13, Lake Illawarra. x 55 Spiral view; 6, Ammobaculites sp. Sample Y10, Lake Illawarra. x50 Spiral view; 7-8, Haplophragmoides canariensis Sample Y52, Windang Island. x47. Umbilical and apertural views; 9, 16, Gaudryina convexa Sample Y52, Windang Island. x25, x44 Lateral views; 10-11, Protoschista findens Sample 306, Lake Illawarra. x50 Lateral views; 12-13, Reophax sp. Sample Y13, Lake Illawarra. x75 Lateral views; 14-15, Milammina fusca Sample 245, Lake Illawarra. x71 Lateral views; 17-20, Eggerella australis Sample 138, Lake Illawarra. x80 Lateral and apertural views; 21, Zéxtularza sp. 1 Sample Y52, Windang Island. x78 Lateral view; 22, Textularia porrecta Sample Y30, Lake Hlawarra. x50 Lateral view; 23, 27, Textularia canderana Sample Y42, Windang Island. x57, x60 Lateral and apertural views; 24, 25, Textularva sagittula Sample Y52, Windang Island. x38 Apertural and lateral views; 26, Zéxtularia sp. II Sample Y52, Windang Island. x48 Lateral view. PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 250 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 251 — , 1933. — The Foraminifera of the tropical Pacific collections of the Albatross, 1899-1900. Bull. U.S. natn. Mus. 161, Pt 2. —— _, 1942. — The Foraminifera of the tropical Pacific collections of the Albatross, 1899-1900. Bull. U.S. natn. Mus. 161, Pt 4. ELIor, I. G., YOUNG, R. W., and CLARKE, D. J., 1976. — Lake hydrology. An Environmental Assessment Project, City of Wollongong Council and University of Wollongong, 41-50. ELLIS, J.. and KANAMORI, S., 1977. — Water pollution studies on Lake Hlawarra. III Distribution of heavy metals in sediments. Aust. J. mar. Freshwat. Res. 28: 485-96. 5 , and LairD, P. G., 1977. — Water pollution studies on Lake Hlawarra. I Salinity variation and estimation of residence time. Aust. J. mar. Freshwat. Res. 30: 467-77. GBs, P. J., 1986. — Five N.S.W. barrier lagoons; their macrobenthic fauna and seagrass communities. Ken- sington, N.S.W.: University of New South Wales, Ph.D. thesis, unpubl. Harris, M., 1976a. — Ecological milieu of Lake Hlawarra. An Environmental Assessment Project, City of Wollon- gong Council and University of Wollongong, 51-65. ——., 1976b. — The biota of Ilawarra Lake. An Environmental Assessment Project, City of Wollongong Council and University of Wollongong, 66-76. Harris, J., 1977. — Ecological studies on Hlawarra Lake with special reference to Zostera capricorni Ascherson. Kensington, N.S.W.: University of New South Wales, M.Sc. thesis, unpubl. HEAN, D., and NANSON, G., 1985. — Geomorphological investigation of stream channels and alluvial deposits in areas of proposed urban development within the catchment of Lake Ilawarra. Report for New South Wales State Pollution Control Commission, Uniadvice, University of Wollongong. HERMELIN, J. O. R., and Scort, D. B., 1985. — Recent benthonic Foraminifera from the north central At- lantic. Micropaleontology 31: 199-220. Jones, B. G., ELior, I. G., and DEPERS, A. M., 1976. — Sediments in Lake Illawarra. An Environmental Assess- ment Project, City of Wollongong Council and University of Wollongong, 20-40. KOHL, B., 1985. — Early Pliocene benthonic foraminifers from the Salina Basin, southeastern Mexico. Bull. Am. Paleont. 88: 1-173. LOEBLICH, A. R., and Tappan, H., 1964. — Treatise on Invertebrate Paleontology. Part C: Protista, Vols 1-2. Lawrence, Kansas: Univ. Kansas Press. , and , 1974. — Recent advances in the classification of the Foraminiferida. Foraminifera 1: 1-53. , and ——., 1984. — Suprageneric classification of the Foraminiferida (Protozoa). Micropaleontology 30: 1-70. Matosa, Y., 1970. — Distribution of Recent shallow-water Foraminifera of Matsushima Bay, Miyagi Prefec- ture, northwest Japan. Scz. Rep. Tohoku Univ., Series 2 (Geol.) 42: 1-85. McCuLLocH, I., 1977. — Quantztative observations on Recent foraminiferal tests with emphasis on the eastern Pacific, Pts 1-3. University of Southern California Publication. ——.,, 1981. — Quantitative observations on Recent foraminiferal tests with emphasis on the eastern Pacific, Pt 4. Univer- sity of Southern California Publication. MIcHIE, M. G., 1982. — Use of the Bray-Curtis similarity measure in cluster analysis of foraminiferal data. J. Math. Geol. 14: 661-667. Murray, J. W., 1973. — Distribution and Ecology of Living Benthonic Foraminiferids. London: Heinemann Educational Books. , 1975. — An Allas of British Recent Foraminiferrds. London: Heinemann Educational Books. Parr, W. J., 1932. — Victorian and South Australian shallow water Foraminifera: Part 2. Proc. Roy. Soc. Vict. 44: 218-234. ——., 1945. — Recent Foraminifera from Barwon Heads, Victoria. Proc. Roy. Soc. Vict. 56: 189-218. ——., 1950. — Foraminifera. British and New Zealand Antarctic Research Expedition 1929-1931 Report Series B, 5: 23323925 PoaG, C. W., 1981. — Ecologic Atlas of Benthic Foraminifera of the Gulf of Mexico. Hutchinson Ross Publ., 174 pp. PUBLIC WORKS DEPARTMENT, N.S.W., 1985. — Lake Illawarra Entrance Study, 109 pp. Fig. 11. 1-3, Eggerella subconica Sample Y 41, Windang Island. x53 Umbilical, spiral and apertural views; 4-6, Tntaxis conica Sample 282, tidal channel. x100 Apertural, lateral and spiral views; 7, Ammotiwm cassis Sample Y30, Lake Illawarra. x56 Lateral view; 8-10, Triloculina oblonga Sample Y33, Lake Illawarra. x70 Spiral and lateral views; 11-12, Quinqueloculina poeyana Sample Y41, Windang Island. x57 Spiral and apertural views; 13, 20, Spiroloculina antillarum Sample Y52, Windang Island. x31 Spiral views; 14-16, Mzlvolinella circulars Sample Y42, Windang Island. x70 Spiral and apertural views; 17-18, Quinqueloculina tropicalis Sample Y 41, Windang Island. x70 Spiral and apertural views; 19, Spiroloculina communis Sample Y52, Windang Island. x36 Spiral view; 21-22, Quinqueloculina sp. nov. Sample Y52, Windang Island. x47 Spiral and apercural views; 23, Quin- queloculina pseudoreticulata Sample Y19, tidal channel. x18 Apertural view; 24-25, Triloculina trigonula Sample Y52, Windang Island. x61 Spiral and apertural views; 26, Miliolinella circularis subsp. nov. Sample Y52, Windang Island. x43 Spiral view. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 OMMUNITIES IN LAKE ILLAWARRA Y 4 FORAMINIFERAL C I. YASSINI AND B. G. JONES 253 Qultty, P. G., 1976. — Foraminifera of Hardy Inlet, southwestern Australia. /. Proc. Roy. Soc. West. Aust. 59: 79-90. Roy, P.S., and Peat, C., 1975. — Bathymetry and bottom sediments of Lake Illawarra. Rec. geol. Surv. N.S.W. WOo=79), SEIBOLD, I., 1975. — Benthonic Foraminifera from the coast and lagoon of Cochin (South India). Revista espa- nola Micropaleontologia 7: 175-213. SIDEBOTTOM, H., 1912. — Lagenae of the south-west Pacific Ocean, from soundings taken by H.M.S. Water- witch 1895. J. Quekett microsc. Club, ser. 2 70: 375-437. ——, 1913. — Lagenae of the south-west Pacific Ocean, from soundings taken by H.M.S. Waterwitch 1895. J. Quekett microsc. Club, ser. 2, Supplementary Paper 12, 73: 161-210. YASSINI, I., and JONES, B. G., 1987. — Ostracoda in Lake Ilawarra: environmental factors, assemblages and systematics. Aust. J. mar. Freshwat. Res., 38: 795-843. , and WRIGHT, A. J., 1988. — Distribution and ecology of Recent ostracodes from Port Hacking, New South Wales. Proc. Linn. Soc. N.S.W. 10: 159-174. Fig. 12. 1-3, Quinqueloculina subpolygona Sample Y52, Windang Island. Details of ornamentation (x460), spiral views (x31, x54); 4-6, Quinqueloculina seminula Sample 282, tidal channel. Spiral and apertural views (x38), details of ornamentation (x460); 7-8, Quinqueloculina granulocostata Sample Y 41, Windang Island. Spiral view (x43), details of ornamentation (x460); 9-14, Milolinella baragwanathi Sample Y52, Windang Island. Lateral, spiral and apertural views (x72), details of ornamentation (x460); 15, Pyrgo subglobulus Sample Y19, tidal channel. x40 Apertural view; 16, Quinqueloculina pseudoreticulata Sample Y19, tidal channel. x40 Apertural view; 17, Sigmoidella elegantissima Sample Y19, tidal channel. x32 Spiral view; 18, Guttulina pacifica Sample Y19, tidal channel. x72 Spiral view; 19, Guttulina sp. 1 Sample Y42, Windang Island. x48 Spiral view; 20, Oolina lineata Sample Y19, tidal channel. x93 Lateral view; 21, Lagena acuticosta Sample Y52, Windang Island. x91 Lateral view; 22, Lagena gracillima Sample Y52, Windang Island. x79 Lateral view; 23, Guttulzna sp. I Sample Y19, tidal channel. x20 Lateral view. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 nN on Ss FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 25 Ol APPENDIX SPECIES IDENTIFIED FROM THE LAKE ILLAWARRA AREA Characteristic features and surface ornamentation of foraminifers from the Lake Illawarra lagoonal, tidal channel and adjacent intertidal environments are listed and illustrated following the classification of Loeblich and Tappan (1964, 1974, 1984). The species are listed in alphabetical order within genera and fami- lies and the scanning electron microscope figures show the characteristic features used for their identification. Family: Rzehakinidae Cushman, 1933 Genus: Miliammina Heron-Allen & Earland, 1930 Militammina fusca (Brady, 1870) CPC 27318 (Fig. 10, nos 14-15) Family: Hormosinidae Haeckel, 1894 Genus: Protoschista Eimer & Fickert, 1899 Protoschista findens (Parker, 1870) CPC 27319 (Fig. 10, nos 10-11) Genus: Reophax Montfort, 1808 Reophax sp. CPC 27320 (Fig. 10, nos 12-13) Family: Lituolidae de Blainville, 1825 Genus: Ammobaculites Cushman, 1910 Ammobaculites agglutinans (d’Orbigny, 1846) CPC 27321 (Fig. 10, no. 5) Ammobaculites foliaceus (Brady, 1881) CPC 27322 (Fig. 10, no. 4) Ammobaculites sp. (Fig. 10, no. 6) Genus: Ammotium Loeblich & Tappan, 1953 Amnotium cassis (Parker, 1870) CPC 27323 (Fig. 11, no. 7) Genus: Haplophragmoides Cushman, 1910 Haplophragmoides canariensis (d’Orbigny, 1839) CPC 27324 (Fig. 10, nos 7-8) Family: Trochamminidae Schwager, 1877 Genus: Tritaxis Schubert, 1921 Tritaxis conica (Parker & Jones, 1865) CPC 27325 (Fig. 11, nos 4-6) Genus: Trochammina Parker & Jones, 1859 Trochammina inflata (Montagu, 1808) CPC 27326 (Fig. 10, nos 1-3) Family: Textularitidae Ehrenberg, 1838 Genus: Jextularia Defrance, in de Blainville, 1824 Textularia candeiana (d’Orbigny, 1839) CPC 27327 (Fig. 10, nos 23, 27) Textularia porrecta (Brady, 1884) CPC 27328 (Fig. 10, no. 22) Textularia sagittula (Defrance, 1824) CPC 27329 (Fig. 10, nos 24-25) Textularia sp. 1 CPC 27330 (Fig. 10, no. 21) Textularia sp. I1 (Fig. 10, no. 26) Fig. 13. 1, Brizalina alata Sample 282, tidal channel. x65 Lateral view; 2, Bolivina doniezi Sample Y19, tidal channel. x70 Lateral view; 3, Brizalina striatula Sample 282, tidal channel. x71 Lateral view; 4, Bolivina robusta Sample Y19, tidal channel. x117 Lateral view; 5, Uvigerina bassensis Sample 282, tidal channel. x67 Lateral view; 6, Bolivina perforatum Sample Y19, tidal channel. x280 Lateral view; 7, Buliminoides gracilis Sample Y19, tidal channel. x72 Lateral view; 8, Buliminoides williamsonianus Sample Y19, tidal channel. x62 Lateral view; 9-10, Bolivina pseudoplicata Sample Y19, tidal channel. x78, x85 Lateral view; 11, Bolzvina sp. Sample 282, tidal channel. xl17 Lateral view; 12, Reussella spinulosa Sample Y19, tidal channel. x70 Lateral view; 13, Lagena crenata Sample 282, tidal channel. x86 Lateral view; 14, Lagena semilineata Sample Y19, tidal channel. x122 Lateral view; 15, Lagena cf. implicata Sample Y19, tidal channel. x68 Lateral view; 16, Planularia patens Sample 282, tidal channel. x72 Lateral view; 17, Bulimina elongata subulata Sample 282, tidal channel. x126 Lateral view; 18, Bulimina marginata Sample Y19, tidal channel. x84 Lateral-apertural view; 19,20, Oolina striatopunc- tata gemma Sample Y19, tidal channel. x193, x79 Lateral views; 21, Fissurina marginatoperforata Sample Y19, tidal channel. x118 Lateral view; 22, Fissurina sulcata Sample Y19, tidal channel. x123 Lateral view; 23, Lagena cf. gracillima Sample Y19, tidal channel. x78 Lateral view; 24, Fissurzna sp. I Sample 282, tidal channel. x100 Lateral view. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA 256 AW ne ’ be ry; Nh — = LIN} PROC I. YASSINI AND B. G. JONES 2S) Family: Ataxophragmiidae Schwager, 1877 Genus: Eggerella Cushman, 1933 Eggerella australis (Collins, 1958) CPC 27331 (Fig. 10, nos 17-20) Eggerella subconica (Parr, 1950) CPC 27332 (Fig. 11, nos 1-3) Genus: Gaudryina d’Orbigny, in de la Sagra, 1839 Gaudryina convexa (Karrer, 1865) CPC 27333 (Fig. 10, nos 9, 16) Family: Nubeculariidae Jones, 1875 Genus: Spivoloculina dOrbigny, 1826 Spiroloculina antillarum (VOrbigny, 1839) CPC 27334 & 27335 (Fig. 11, no. 13) Spiroloculina communis (Cushman & Todd, 1944) (Fig. 11, no. 19) Family: Miliolidae Ehrenberg, 1839 Genus: Miliolinella Wiesner, 1931 Miliolinella baragwanathi (Parr, 1945) CPC 27337 (Fig. 12, nos 9-14) Miliolinella circularis (Bornemann, 1855) CPC 27338 (Fig. 11, nos 14-16) Milolinella circulars subsp. nov. CPC 27339 (Fig. 11, no. 26) Miliolinella subrotunda (Montagu, 1803) (Fig. 19, nos 1-4) Genus: Pyrgo Defrance, in de Blainville, 1824 Pyrgo subglobulus (Parr, 1950) CPC 27340 (Fig. 12, no. 15) Genus: Quinqueloculina dOrbigny, 1826 Quinqueloculina granulocostata (Germeraad, 1946) CPC 27341 (Fig. 12, nos 7-8) Quinqueloculina poeyana (d’Orbigny, 1839) CPC 27342 (Fig. 11, nos 11-12) Quinqueloculina pseudoreticulata (Parr, 1941) CPC 27343 (Fig. 11, no. 23, Fig. 12, no. 16) Quinqueloculina seminula (Linné, 1767) CPC 27344 (Fig. 12, nos 4-6) Quinqueloculina subpolygona (Parr, 1945) CPC 27345 (Fig. 12, nos 1-3) Quinqueloculina tasmanica (Albani, 1978) CPC 27346 (Fig. 11, no. 20) Quinqueloculina tropicalis (Cushman, 1924) CPC 27347 (Fig. 11, nos 17-18) Quinqueloculina sp. nov. CPC 27348 (Fig. 11, nos 21-22) Genus: Triloculina d’Orbigny, 1826 Triloculina oblonga (Montagu, 1803) CPC 27350 & 27351 (Fig. 11, no. 8-10) Triloculina tricarinata (dOrbigny, 1826) Triloculina trigonula (Lamarck, 1804) CPC 27352 (Fig. 11, nos 24-25) Family: Polymorphinidae d’Orbigny, 1839 Genus: Guttulina d’Orbigny, in de la Sagra, 1839 Guttulina pacifica (Cushman & Ozawa, 1928) CPC 27355 (Fig. 12, no. 18) Guttulina regina (Brady, Jones & Parker, 1870) CPC 27356 (Fig. 19, no. 5) Guttulina sp. 1 CPC 27357 (Fig. 12, no. 19) Guttulina sp. UW CPC 27358 (Fig. 12, no. 23) Genus: Sigmozdella Cushman & Ozawa, 1928 Sigmoidella elegantissima (Parker & Jones, 1865) CPC 27359 (Fig. 12, no. 17) Family: Nodosariidae Ehrenberg, 1838 Genus: Amphicoryna Schlumberger, in Milne-Edward, 1881 Amphicoryna scalaris (Batsch, 1791) CPC 27360 (Fig. 15, no. 10) Fig. 14. 1, Lagena subacuticosta Sample 282, tidal channel. x83 Lateral view; 2, Lagena sulcata Sample 282, tidal channel. x117 Lateral view; 3, Fissurina lacunata Sample Y19, tidal channel. x108 Lateral view; 4, Fissurina fasciata carinata Sample Y52, Windang Island. x102 Lateral view; 5, Fissurina cf. subquadrata Sample Y52, Windang Island. x118 Lateral view; 6, Lagena cf. semistriata Sample Y19, tidal channel. x67 Lateral view; 7, Oolina sp. | Sample Y19, tidal channel. x117 Lateral view; 8, Oolina sp. I] Sample 282, tidal channel. x101 Lateral view; 9, Fissurina sp. II Sample Y19, tidal channel. x83 Lateral view; 10, Bolivinella folia Sample Y19, tidal channel. x77 Lateral view; 11, Lamarckina sp. Sample Y19, tidal channel. x71 Spiral view; 12, Lamarckina sp. Sample Y42, Windang Island. x71 Umbilical view; 13-14, Patellina corrugata Sample Y52, Windang Island. x87 Spiral and umbilical views; 15-16, Glabratella tabernacularis Sample Y42, Windang Island. x91 Spiral and umbilical views; 17-20, Glabratella australensis Sample Y52, Windang Island. x60, x91 Spiral, plastogamic and umbilical views; 21-22, Glabratella sp. 1. Sample Y52, Windang Island. x70 Umbilical and spiral views. PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 258 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 259 Genus: Bolivinella Cushman, 1927 Bolivinella folia (Parker & Jones, 1865) CPC 27361 (Fig. 14, no. 10) Genus: Fissurina Reuss, 1850 Fissurina fasciata carinata (Sidebottom, 1906) CPC 27362 (Fig. 14, no. 4) Fissurina lacunata (Burrows & Holland, 1895) CPC 27363 (Fig. 14, no. 3) Fissurina marginatoperforata (Seguenza, 1913) CPC 27364 (Fig. 13, no. 21) Fissurina sp. cf. subquadrata (Parr, 1945) CPC 27365 (Fig. 14, no. 5) Fissurina sulcata (Collins, 1974) CPC 27366 (Fig. 13, no. 22) Fissurina sp. | CPC 27367 (Fig. 13, no. 24) Fissurina sp. 11 CPC 27368 (Fig. 14, no. 9) Genus: Lagena Walker & Jacob, in Kanmacher, 1798 Lagena acuticosta (Reuss, 1861) CPC 27369 (Fig. 12, no. 21) Lagena crenata (Parker & Jones, 1865) (Fig. 13, no. 13) Lagena gracillima (Seguenza, 1862) CPC 27370 (Fig. 12, no. 22) Lagena sp. cf. gracillima (Seguenza, 1862) CPC 27371 (Fig. 13, no. 23) Lagena sp. cf. implicata (Cushman & McCulloch, 1950) CPC 27372 (Fig. 13, no. 15) Lagena semilineata (Wright, 1866) CPC 27373 (Fig. 13, no. 14) Lagena sp. cf. semistriata (Williamson, 1848) CPC 27374 (Fig. 14, no. 6) Lagena subacuticosta (Parr, 1950) (Fig. 14, no. 1) Lagena sulcata (Walker & Jacob, 1798) CPC 27375 (Fig. 14, no. 2) Genus: Planularia Defrance, in de Blainville, 1824 Planularia patens (Brady, 1884) CPC 27376 (Fig. 13, no. 16) Family: Glandulinidae Reuss, 1860 Genus: Oolina d’Orbigny, in de la Sagra, 1839 Oolina lineata (Williamson, 1848) CPC 27377 (Fig. 12, no. 20) Oolina striatopunctata gemma (Cushman & McCulloch, 1950) CPC 27378 (Fig. 13, nos 19-20) Oolina sp. 1 CPC 27379 (Fig. 14, no. 7) Oolina sp. Il CPC 27380 (Fig. 14, no. 8) Family: Bolivinitidae Cushman, 1927 Genus: Bolivina d’Orbigny, in de la Sagra, 1839 Bolivina compacta (Sidebottom, 1905) CPC 27381 Bolivina doniezi (Cushman & Wickenden, 1928) (Fig. 13, no. 2) Bolivina perforatum (Dinapoli, 1946) (Fig. 13, no. 6) Bolivina pseudoplicata (Heron-Allen & Earland, 1930) CPC 27382 (Fig. 13, nos 9-10) Bolivina robusta (Brady, 1881) CPC 27383 (Fig. 13, no. 4) Bolivina sp. CPC 27384 (Fig. 13, no. 11) Genus: Brizalina Costa, 1856 Brizalina alata (Seguenza, 1862) CPC 27385 (Fig. 13, no. 1) Brizalina striatula (Cushman, 1922) CPC 27386 (Fig. 13, no. 3) Family: Buliminidae Jones, 1875 Genus: Bulimina d’Orbigny, 1826 Bulimina elongata subulata (Cushman & Parker, 1937) (Fig. 13, no. 17) Bulimina marginata (d’Orbigny, 1826) CPC 27388 (Fig. 13, no. 18) Genus: Reussella Galloway, 1933 Reussella spinulosa (Reuss, 1850) CPC 27389 (Fig. 13, no. 12) Fig. 15. 1, Uvigerina cf. peregrina Sample Y19, tidal channel. x60 Lateral view; 2-3, Angulodiscorbis quadrangularis Sample Y52, Windang Island. x93 Lateral and umbilical views; 4, Patellinella inconspicua Sample Y42, Windang Island. x105 Umbilical and lateral views; 5-7, Buccella pustulosa Sample Y52, Windang Island. x115 Apertural, umbilical and spiral views; 8-9, Rosalina anglica Sample Y52, Windang Island. x48 Spiral and umbilical views; 10, Amphicoryna scalaris Sample Y19, tidal channel. x60 Lateral view; 11-14, Rosalina bradyi Sample Y52, Windang Island. x63 Umbilical, spiral, apertural and floating chamber views; 15, Rosalina australis Sample Y52, Windang Island. x65 Spiral view; 16-18, Glabratella pulvinata Sample Y52, Windang Island. x93 Spiral, lateral and umbilical views; 19-20, Cymbaloporetta bradyi Sample Y52, Windang Island. x43 Umbilical and spiral views; 21-23, Glabratella sp. II Sample Y19, tidal channel. x86 Spiral and umbilical views; 24-25, Planulinoides biconcavus Sample 282, tidal channel. x65 Spiral and apertural views. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 260 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA Fig. 16. 1-3, Elphidium depressulum Sample Y52, Windang Island. Details of lateral process (x460), umbilical and apertural views (x80); 4-5, Elphidium advenum Sample Y42, Windang Island. Details of lateral process (x460), and umbilical view (x87); 6-8, Elphidium macellum Sample Y42, Windang Island. Details of lateral process (x460), and spiral view (x60); 9-11, Cribrononion sydneyensis Sample 390, Lake Illawarra. Details of lateral process (x460), and spiral and apertural views (x50); 12-14, Elphidium argenteus Sample Y52, Windang Island. x52 Lateral-apertural and umbilical views; 15-16, Nontonella auris Sample Y19, tidal channel. x72 Lateral-apertural and umbilical views; 17-18, Nonion depressulus Sample Y19, tidal channel. x87 Umbilical and apertural views PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 261 Family: Turrilinidae Cushman, 1927 Genus: Buliminoides Cushman, 1911 Buliminoides gracilis (Collins, 1953) CPC 27391 (Fig. 13, no. 7) Buliminoides williamsonianus (Brady, 1881) CPC 27392 (Fig. 13, no. 8) Family: Uvigerinidae Haeckel, 1894 Genus: Uvigerina dOrbigny, 1826 Uvigerina bassensis (Parr, 1950) CPC 27395 (Fig. 13, no. 5) Uvigerina sp. cf. peregrina (Cushman, 1923) (Fig. 15, no. 1) Family: Glabratellidae Loeblich & Tappan, 1963 Genus: Angulodiscorbis Uchio, 1953 Angulodiscorbis quadrangularis (Uchio, 1953) CPC 27396 (Fig. 15, nos 2-3) Genus: Glabratella Dorreen, 1948 Glabratella australensis (Heron-Allen & Earland, 1932) CPC 27397 (Fig. 14, nos 17-20) Glabratella sp. cf. parrt (Collins, 1974) Glabratella patelliformis (Brady, 1884) CPC 27398 Glabratella pulvinata (Brady, 1884) CPC 27399 (Fig. 15, nos 16-18) Glabratella tabernacularis (Brady, 1884) CPC 27400 (Fig. 14, nos 15-16) Glabratella sp. 1 CPC 27401 (Fig. 14, nos 21-22) Glabratella sp. 11 CPC 27402 (Fig. 15, nos 21-23) Family: Discorbidae Ehrenberg, 1838 Genus: Baggina Cushman, 1926 Baggina phillippinensis (Cushman, 1921) CPC 27403 (Fig. 17, nos 18-19) Genus: Buccella Andersen, 1952 Buccella pustulosa (Albani, 1982) CPC 27404 (Fig. 15, nos 5-7) Genus: Discorbinella Bandy, 1949 Discorbinella berthelots (d’Orbigny, 1839) CPC 27405 (Fig. 18, nos 13-14) Discorbinella planoconcava (Chapman, Parr & Collins, 1932) CPC 27406 (Fig. 18, no. 25) Genus: Lamellodiscorbis Bermudez, 1952 Lamellodiscorbis dimidiatus (Jones & Parker, 1862) CPC 27407 (Fig. 17,nos 9-11) Genus: Patellinella Cushman, 1928 Patellinella inconspicua (Brady, 1884) CPC 27408 (Fig. 15, no. 4) Genus: Planulinoides Parr, 1941 Planulinoides biconcavus (Jones & Parker, 1862) CPC 27409 (Fig. 15, nos 24-25) Family: Rosalinidae Reiss, 1963 Genus: Rosalina dOrbigny, 1826 Rosalina anglica (Cushman, 1931) CPC 27410 (Fig. 15, nos 8-9) Rosalina australis (Parr, 1932) CPC 27411 (Fig. 15, no. 15) Rosalina brady: (Cushman, 1951) CPC 27412 (Fig. 15, nos 11-14) Family: Robertinidae Reuss, 1850 Genus: Lamarckina Berthelin, 1881 Lamarckina sp. CPC 27413 (Fig. 14, nos 11-12) Family: Cymbaloporidae Cushman, 1927 Genus: Cymbaloporetta Cushamn, 1928 Cymbaloporetta bradyi (Cushman, 1915) CPC 27414 & 27415 (Fig. 15, nos 19-20) Family: Planorbulinidae Schwager, 1877 Genus: Acervulina Schultz, 1854 Acervulina inhaerens (Schultze, 1854) CPC 27416 & 27417 (Fig. 17, nos 14-15) Genus: Planorbulina @Orbigny, 1826 Planorbulina mediterranensis (d’Orbigny, 1826) CPC 27418 (Fig. 17, no. 12) PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 262 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA Fig. 17.1-2, Parrellina imperatrix Sample Y42, Windang Island. Details of lateral process (x460), and spiral view (x20); 3-4, Elphidium sp. Sample Y19, tidal channel. Details of lateral process (x460), and spiral view (x91); 5-6, Elphidium crispum Sample 282, tidal channel. Details of lateral process (x460), and spiral view (x31); 7-8, Elphidium jenseni Sample 282, tidal channel. Details of lateral process (x460), and spiral view (x20); 9-11, Lamellodiscorbis dimidiatus Sample Y52, Windang Island. x26 Umbilical, lateral-apertural and spiral views; 12, Planorbulina mediterranensis Sample Y18, tidal channel. x27 Spiral view; 13, Spirillina vivipara Sample 282, tidal channel. x86 Spiral view; 14-15, Acervulina inhaerens Sample Y52, Windang Island. x63 Umbilical and piral views; 16, Spirillina tuberculata Sample Y19, tidal channel. x55 Umbilical view; 17, Spirillina sp. Sample Y19, tidal channel. x95 Umbilical view; 18-19, Baggina phillippinensis Sample Y52, Windang Island. x52 Spiral and umbilical views. PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES Family: Elphidiidae Galloway, 1933 Genus: Cribrononion Thalmann, 1947 Cribrononion sydneyensis (Albani, 1978) CPC 27424 (Fig. 16, nos 9-11) Genus: Elphidium Montfort, 1808 Elphidium advenum (Cushman, 1922) CPC 27425 (Fig. 16, nos 4-5) Elphidium argenteus (Parr, 1945) CPC 27426 (Fig. 16, nos 12-14) Elphidium crispum (Linne, 1758) CPC 27427 (Fig. 17, nos 5-6) Elphidium depressulum (Cushman, 1933) (Fig. 16, nos 1-3) Elphidium jensent (Cushman, 1924) CPC 27428 (Fig. 17, nos 7-8) Elphidium macellum (Fichtel & Moll, 1798) CPC 27429 (Fig. 16, nos 6-8) Elphidium sp. CPC 27430 (Fig. 17, nos 3-4) Genus: Parrellina Phalmann, 1951 Parrellina imperatrix (Brady, 1881) CPC 27431 (Fig. 17, nos 1-2) Family: Nonionidae Schultze, 1854 Genus: Nonion Montfort, 1808 Nonton depressulus (Walker & Jacob, 1798) (Fig. 16, nos 17-18) Genus: Nonionella Cushman, 1926 Nonvonella aurs (d’Orbigny, 1839) CPC 27434 (Fig. 16, nos 15-16) Family: Rotaliidae Ehrenberg, 1839 Genus: Ammonia Brunnich, 1772 Ammonia beccari (Linné, 1767) CPC 27432 (Fig. 18, no. 8) Genus: Rotalia Lamarck, 1804 Rotalia perlucida (Heron-Allen & Earland, 1913) CPC 27433 (Fig. 18, nos 5-7) Family: Cibicididae Cushman, 1927 Genus: Crbicides Montfort, 1808 Cibicides cygnorum (Carter, 1964) CPC 27419 (Fig. 18, nos 1-2, 12) Cibicides refulgens (Montfort, 1808) CPC 27420 Genus: Dyoczbicides Cushman & Valentine, 1930 Dyocibicides biserralts (Cushman & Valentine, 1930) CPC 27421 (Fig. 18, nos 9-10) Family: Anomalinidae Cushman, 1927 Genus: Anomalina dOrbigny, 1826 Anomalina nonionordes (Parr, 1932) CPC 27435 (Fig. 18, no. 11) Family: Spirillinidae Reuss, 1862 Genus: Patellina Williamson, 1858 Fatellina corrugata (Williamson, 1858) CPC 27436 (Fig. 14, nos 13-14) Genus: Spirillina Ehrenberg, 1843 Spirillina denticulata (Brady, 1884) CPC 27437 (Fig. 18, nos 3-4) Spirillina inaequalis (Brady, 1879) CPC 27438 Spirillina tuberculata (Brady, 1884) CPC 27439 (Fig. 17, no. 16) Spirllina vivipara (Ehrenberg, 1843) CPC 27440 (Fig. 17, no. 13) Spirillina sp. CPC 27441 (Fig. 17, no. 17) Family: Globigerinidae d’Orbigny, 1826 Genus: Globigerina d’Orbigny, 1826 Globigerina bulloides (d’Orbigny, 1826) CPC 27442 (Fig. 18, no. 15) Globigerina sp. Genus: Globigerinoides Cushman, 1927 Globigerinoides ruber (d’Orbigny, 1839) CPC 27443 (Fig. 18, nos 19-20) Genus: Globoquadrina Finlay, 1947 Globoquadrina dutertre: (d’Orbigny, 1839) CPC 27444 (Fig. 18, nos 21-22) Genus: Orbulina @Orbigny, in de la Sagra, 1839 Orbulina universa (d’Orbigny, 1839) CPC 27445 (Fig. 18, no. 16) Genus: Pulleniatina Cushman, 1927 Pulleniatina obliqueloculata (Parker & Jones, 1865) 263 PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 264 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA xt SNR oes eo PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 I. YASSINI AND B. G. JONES 265 Family: Globorotaliidae Cushman, 1927 Genus: Globorotalia Cushman, 1927 Globorotalia hirsuta (dOrbigny, 1839) CPC 27446 (Fig. 18, nos 23-24) Subgenus: TZurborotalia Cushman & Bermudez, 1949 Globorotalia ( Turborotalia) inflata (d’Orbigny, 1839) CPC 27447 (Fig. 18, nos 17-18) Fig. 18. 1-2, 12, Crbicrdes cygnorum Sample 282, tidal channel. x78 Umbilical, spiral and apertural views; 3-4, Spirillina denticulata Sample Y19, tidal channel. Details of ornamentation (x460), and spiral view (x74); 5-7, Rotalia perlucida Sample Y52, Windang Island. x70 Apertural, spiral and umbilical views; 8, Ammonia beccarit Sample Y10, Lake Illawarra. x43 Umbilical view; 9-10, Dyocibicides biserialis Sample Y42, Windang Island. x43 Umbilical and spiral views; 11, Anomalina nonionoides Sample Y19, tidal channel. x50 Spiral view; 13-14, Discorbinella bertheloti Sample Y19, tidal channel. x87 Spiral and umbilical views; 15, Globigerina bullovdes Sample Y19, tidal channel. x71 Umbilical view; 16, Orbulina universa Sample Y19, tidal channel. x64; 17-18, Globorotalia (Turborotalia) inflata Sample 282, tidal channel. x71 Umbilical and apertural views; 19-20, Globigerinoides ruber Sample Y19, tidal channel. x72 Spiral, lateral-apertural views; 21-22, Globoquadrina dutertre: Sample Y19, tidal channel. x70 Umbilical and spiral views; 23-24, Globorotalia hirsuta Sample Y19, tidal channel. x70 Umbilical and spiral views; 25, Discorbinella planoconcava Sample Y42, Windang Island. x90 Umbilical view. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 266 FORAMINIFERAL COMMUNITIES IN LAKE ILLAWARRA Fig. 19. 1-4, Miliolinella subrotunda Sample Y28, Lake Illawarra. 1, 4 apertural views (x200); 2 spiral view (x170); 3 lateral view (x160); 5, Guttulina regina Sample Y42, Windang Island. x60 Lateral view. PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 Status of the Genera Ophiopeza and Ophiopsammus (Echinodermata: Ophiuroidea) in Australian Waters, with the Description of a new Species Ee. VAtiand F WoE ROWE VaIL, L. L., & Rowe, F. W. E. Status of the genera Ophiopeza and Ophiopsammus (Echinodermata: Ophiuroidea) in Australian waters, with the description of a new species. Proc. Linn. Soc. N.S.W. 110(3), (1988) 1989: 267-288. A study of type and other material has resulted in a reappraisal of the generic limits of the ophiodermatid genera Ophiopeza Peters, 1851 and Ophiopsammus Lutken, 1869. By using disc scaling as a character of generic significance the species fallax arabica A. M. Clark, 1968, cylindrica Hutton, 1872a, and spinosa, Ljungman, 1867, have been retained in the genus Ophiopeza, type-species fallax fallax Peters, 1851, while the species aequalis Lyman, 1880, anchista H. L. Clark, 1911, assimilis Bell, 1888, and maculata Verrill, 1869 have been transferred to the genus Ophiopsammus, type-species yoldi: Lutken, 1856. In addition, Ophiopsammus angusta sp. nov. 1s described from southeastern Australian waters. The following species have been synonymized: Ophiopeza arenosa (Lyman, 1879) and Ophiopeza gracilis (Mortensen, 1924) with Ophiopeza cylindrica; Ophiopeza dubiosa (de Loriol, 1893) with Ophiopeza spinosa; and Ophiopeza dyscrita (H. L. Clark, 1909) and Ophiopeza nigra (H. L. Clark, 1938) with Ophiopsammus assimilis. Ophiopsammus aequalls 1s recorded for the first time from Australia. Notes are given for a misidentified type speci- men of anchista. A key to all the known species of Ophiopeza and Ophiopsammus is given, except for exzlzs Koehler, 1905. L. L. Vail, Northern Territory Museum, G. PO. Box 4646, Darwin, Australia 5794, and F. W. E. Rowe, Division of Invertebrate Zoology (Echinoderms), Australian Museum, PO. Box A285, Sydney South, Australia 2000; manuscript rececved 30 December 1986, accepted for publication 18 May 1988. INTRODUCTION The taxonomic status of Pectinura Forbes 1843, Ophiopeza Peters 1851, and Ophio- psammus Lutken 1869 has been most recently discussed by A. M. Clark (1968). Briefly, Pectinura was described by Forbes (1843) for vestita from a small specimen taken off southern Turkey. Ophiopeza was described by Peters (1851) for the species fallax which was collected off Mozambique. These two genera were separated on the presence of supplementary oral shields in Pectinura and their absence in Ophiopeza. Ophiopsammus was established by Lutken (1869) for his earlier described species Ophiopeza yoldi. This genus was distinguished by the absence of supplementary oral shields and the concealment of radial shields by disc granules. Lyman (1882) subsequently referred yoldi back to Ophiopeza although he gave no justification. He listed 12 species belonging to the genus Fectinura and five species to Ophiopeza. In a review, H. L. Clark (1909a) synonymized both Ophiopeza and Ophiopsammus with Pectinura on the grounds that the presence or absence of supplementary oral shields in these genera lacked sufficient taxonomic value. However, he stated that if his interpretation of Forbes’ inadequate description of vestita was incorrect then Péctinura should probably replace Ophiarachnella Ljungman, 1871. Furthermore, Ophiopeza would become the available generic name for the 10 species-he © included in the genus F¢ctinura. Later in the same year, he described (H. L. Clark, 1909b) another species of Pectinura, dyscrita, from off New South Wales. During his visits to Aus- tralia in 1913, 1929, and 1932, H. L. Clark collected only one specimen of Pectinura which he described as a new species, nigra (H. L. Clark, 1938) from Western Australia. In his 1946 review of the Australian echinoderm fauna, H. L. Clark recorded five species of PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 268 OPHIOPEZA AND OPHIOPSAMMUS Fectinura from Australian waters, mentioning that none was known from adequate material. Mortensen (1940) recorded Ophiopeza fallax from the Iranian Gulf. He questioned whether the species should be referred to the genus Pectinura but decided that since the type-species of Pectinura, P. vestita, is insufficiently known he preferred to use the name Ophiopeza. A. M. Clark (1968) discussed the problem of these genera and concurred with Mortensen’s use of the name Ophiopeza, especially since the holotype of Pectinura is now lost. She suggested that P vestzta could have been a specimen of the Mediterranean species Ophioconis forbes which had lost disc granules from the oral shields. However, A. M. Clark (1968) did not agree with Lyman’s inclusion of Ophiopsammus in the synonymy of Ophiopeza and, on the basis of the shape of its dorsal arm plates, she revived the generic name Ophiopsammus for the species yoldi1. During the course of researching the echinoderm fauna of New South Wales (FW.E.R.; ARGS D1805325; MST 84/2092) extensive collections made along the coast during 1981 and 1982 by staff of the Echinoderm Section of the Australian Museum yielded a large number of specimens referred to the genus Ophiopeza, as understood by A. M. Clark (1968). This material has been compared with other specimens in the col- lections of the Australian Museum and with type and other specimens borrowed from the British Museum (Natural History), Museum of Comparative Zoology, Museum of Victoria, National Museum of New Zealand, and the US National Museum (Smith- sonian). As a result of our investigations, we have been able to clarify the status of the species of Ophiopeza and Ophiopsammus occurring in Australia, as well as describing a new species and establishing a new record. Furthermore, we have established the value of disc scaling in separating the genera Ophiopeza and Ophiopsammus. With the exception of Lyman’s (1882) use of this feature in a key to the species of Pectinura, disc scaling has been ignored in ophiodermatid taxonomy. The following abbreviations have been used: ca = approximately; coll. = collec- tion; d.d. = disc diameter; d.r. = disc radius; R = arm length from base of disc to arm tip; AM = Australian Museum, Sydney: BM(NH) = British Museum (Natural History), London; MCZ = Museum of Comparative Zoology, Cambridge, Mass.; NMNZ = National Museum of New Zealand, Wellington, MV = Museum of Victoria, Melbourne; USNM = US National Museum (Smithsonian), Washington; N.S.W. = New South Wales; N.Z. = New Zealand: Old = Queensland; S.A. = South Australia; Tas. = Tasmania; Vic. = Victoria; W.A: = Western Australia. All known species in each of the genera (except for Ophiopeza exilis) are included in their respective keys, but only species known from Australian waters are discussed in detail. SYSTEMATIC ACCOUNT Family OPHIODERMATIDAE Genus Ophiopeza Peters Ophiopeza Peters, 1851: 465 (type-species O. fallax Peters, 1851, by monotypy). Ophiopezella Ljungman, 1872: 639 (type-species Ophiarachna spinosa Ljungman, 1867, by monotypy). Fectinura: autorum [non Fectinura Forbes, 1843]. Diagnosis: A genus of ophiodermatid with radial shields usually obscured by granules; marginal disc scales enlarged, usually covered by granules, rarely bare; disc scales coarse, overlapping, 1-3 large scales separating the two plates of each pair of radial shields; oral plates covered by granules; oral shields naked, supplementary oral shields PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 L. L. VAILAND F. W. E. ROWE 269 frequently present and at least partially naked; dorsal arm plates fan shaped to trans- versely rectangular (up to ca 2 x wider than long), flat; arm spines never exceeding one arm segment in length, often appressed. Other species included: O. fallax arabica A. M. Clark, 1968; O. cylindnca (Hutton, 1872a); O. spinosa (Ljungman, 1867). Remarks: We concur with Mortensen (1940) and A. M. Clark (1968) in their recog- nition of the genus Ophiopeza. Also, we are able to support A. M. Clark’s reasoning and decision in her resurrection of the genus Ophiopsammus from synonymy with Ophzopeza. However, in addition, we draw attention to differences in the type of disc scaling between Ophiopeza and Ophiopsammus (see p. 268). Using these characters, we propose to restrict Ophiopeza for fallax fallax, fallax arabica, cylindrica, and spinosa. Of the other species previously referred to Ophiopeza by A. M. Clark (1968), we propose that anchista H. L. Clark, 1911, assemilis Bell, 1888, aequalis Lyman, 1880, and maculata Verrill, 1869, be referred to Ophiopsammus, as we consider them to be congeneric with yoldi Lutken, the type species of Ophiopsammus. We are uncertain of the affinity of exzlzs (Koehler, 1905) which A. M. Clark (1968) transferred back from Pectinura to Ophiopeza as its disc scaling is not evident on the figure given for it by Koehler (1905: pl. 2, fig. 5) and because we have not examined material referable to this species. Ophiopeza cylindrica (Hutton, 1872a) Figs 1A-C, 2A-B Ophiura cylindrica Hutton, 1872a: 3; 1872b: 811. Pectinura arenosa Lyman, 1879: 48, pl. 14, figs 392-394; 1882: 15, pl. 23, figs 10-12. H. L. Clark, 1909: 117; 1921: 141; 1946: 257. A. M. Clark, 1966: 327. Dartnall, 1980: 42, mle Ophiopeza cylindrica. Farquhar, 1895: 198, 1898b: 306 [? non Ophiopeza cylindrica. Farquhar, 1898a: 190, pl. 14, figs 4, 5 = assemilzs.] Pectinura cylindrica. H. L. Clark, 1909: 117; 1915: 303 [? non Pectinura cylindrica. Morten- sen, 1924: 172, non fig. 35(1-2) = assemilss. | Pectinura gracilis Mortensen, 1924: 173, figs 35(3-5), 36. Ophiopeza arenosa. A. M. Clark, 1968: 313. Rowe and Vail, 1982: 222. [? non Ophiopeza arenosa. Baker, 1982: 431, fig. 10.18c. = assimilis. | Material examined: Lectotype of O. cylindrica NUNZ Ech 6, no coll. data; lectotype of O. gracilis NMNZ Ech 375, Paterson Inlet, Stewart Id, N.Z., 9-28m, 17. x1. 14; holotype of O. arenosa BM(NH) 82.12.23.247, off East Mancoeur Id, Bass Strait, Tas., 70-74m, 2.1v.1874; 1 paratype of O. arenosa BM(NH) 82.12.23.247 (pt) off East Mancoeur Id, Bass Strait, Tas., 70-74m, 2.iv.1874; 1 spec., AM G7831, Watson’s Bay, Port Jackson, N.S.W., no coll. depth or date; 1 spec., AM G11413, Broughton Id, nr Port Stephens, N.S.W., 66m, no coll. date; 2 spec., AM J15049, NW Solitary Id, nr Coffs Hbr, N.S.W., 22m, 10.11.82; 1 spec., AM G11032, Port Jackson, N.S.W., no coll. depth; 1 spec., AM J1974, Port Phillip, Vic., no coll. depth; 3 spec., MNMZ (Ech 3412), Paterson Inlet, N.Z., 23m, 26.1.60; 2 spec., NMNZ (ECH 3413), North Arm, Port Pegasus, Stewart Id, N.Z. 37- 44m, 22.11.72. j Diagnosis: Disc diameter to 11mm, densely covered with round granules, 13-16 per mm; coarse, slightly overlapping scales underlying the granules. Each interradius with 7-9 marginal disc plates, middle ones largest, each plate obscured by granule cover. Radial PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 270 OPHIOPEZA AND OPHIOPSAMMUS Fig. 1. A, B, Ophiopeza cylindrica, lectotype (NMNZ Ech 6), dorsal and ventral view respectively, dd = 8mm. C, Ophiopeza gracilis, lectotype (NMNZ Ech 375), doral view, dd = 5mm. Some disc granules were removed in A and C to show disc scaling. shields 0.1 x d.r., obscured by granules. Three conspicuous plates in triangular arrange- ment separating the two plates of each pair of radial shields. Arms to 42mm, 2.6-3.9 x d.d., dorsal plates fan shaped in specimens with d.d. & 7mm becoming transversely rec- tangular in shape (up to 2.0 times wider than long) in larger specimens; ventral plates as long as or slightly longer than wide. Eight to ten short, stout arm spines per segment; lowermost sometimes longer and wider than the others, which were about half an arm segment long. Two tentacle scales per pore, inner one longest, the outer one covering the base of lowest arm spine. Oral shields usually with a distal supplementary plate which is up to % as long as the oral plate; adoral plates roughly triangular in shape. Oral plates covered with granules. Seven to eight short, flat, oral papillae; tooth papillae flattish, rounded distally. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAILAND F. W. E. ROWE 271 Fig. 2. A, B, Ophiopeza arenosa, paratype (BM(NH) 82.12.23.247 (pt) ), dorsal view, dd = 9mm. Synonymized with Ophiopeza cylindrica herein. Some disc granules were removed to show disc scaling. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 DUD OPHIOPEZA AND OPHIOPSAMMUS Colour (dried): Light brown to charcoal grey; some arms with dark, cream, or reddish bands. Distribution: Northeastern Tas. and Port Phillip Bay, Vic. to off Coffs Harbour, N.S.W., Stewart Id, N.Z.; to 74m. Remarks: The description of Ophiura cylindrica by Hutton (1872a) was based on two specimens (Hutton, 1872b); neither of which had locality data according to the original authority. Lyman (1882) synonymized this species with Pectinura rigida Lyman, 1874, although he gave no reasons for the synonymy. Farquhar (1895) compared Lyman’s description of PR rgida with the syntype series of O. cylindrica and concluded that they represented two distinct species. H. L. Clark (1915) subsequently synonymized rgida with Ophiarachnella septemspinosa (Muller and ‘Troschel, 1842). Farquhar (1898a) described and figured material as cylindrica, which we believe refers to Ophiopsammus assimilis on the grounds of dorsal arm plate shape (see below). One of two syntypes of O. cylindrica was already lost when Mortensen (1924) compared this species with his newly- established species, Ophiopeza gracilis. We have examined the remaining syntype speci- men of O. cylindrica (NMNZ Ech 6) examined by Mortensen, and agree with his com- ments (Mortensen, 1924). We are therefore convinced that this is one of Hutton’s syntype series and we designate it as the lectotype of O. cylindrica. O. gracilis (Mortensen, 1924) was established on a number of specimens from New Zealand. These included an unspecified number from Stewart Id, two from Queen Charlotte Sound, and one from Three Kings Island. We designate one specimen from Stewart Id (NMNZ Ech 375) as a lectotype of O. gracilis (Mortensen) since we consider it to be part of his syntype series. When Mortensen described O. gracilis, he compared it with the closely related species cylindrica and also figured each of the species (Mortensen, 1924: 174, fig. 35 (1-2) ). We have material of a size similar to the specimens of cylindrica figured by both Mortensen (1924) and Farquhar (1898a: 190, pl. 14, figs 4, 5). Our material agrees with the specimens figured by both of them although we identify our specimens as Ophio- psammus assimilis on the basis of disc scaling and shape of the dorsal arm plates. Con- sequently, we contend that Mortensen and Farquhar mistakenly referred specimens of assimilis to cylindrica. However, we agree with the differences discussed by Mortensen (1924) between the species ‘cylindrica and gracilis but only in that they refer to differences between Ophiopsammus assimilis (i.e. Mortensen’s and Farquhar’s cylindrica) and cylindrica s.s. (1.e. Mortensen’s gracilis). Mortensen (1924) used dorsal arm plate and arm spine shape, relative size of oral shields, and the extent of granulation on oral plates to differentiate ‘cylindrica’ and gracilis. He concluded that the dorsal arm plates were basically fan shaped in gracilis but that they were more transversely rectangular in ‘cylindrica. Table 1 summarizes measure- ments from 15 specimens we have identified as cylindrica s.s., based primarily on dorsal disc scaling. Dorsal arm plates change from fan shaped in small individuals (d.d. & 7mm) to more transversely rectangular shaped in larger individuals, as shown in the dorsal arm plate ratio column of Table 1. This evidence suggests that the differences referred to by Mortensen can be explained by ontogenic development. Ophiopeza arenosa was established on 6 type specimens by Lyman (1879). We have examined the holotype and a paratype (BM(NH) 82.12.23.247). Type material of arenosa was compared with the lectotype of Ophiopeza cylindrica. In these specimens, only slight differences attributable to size could be detected and these are within the range of vari- ation found in cylindrica (‘Table 1). PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 L. L. VAILANDFE. W. E. ROWE 273 TABLE 1 Measurements of specimens of Ophiopeza cylindrica Registration no. d.d. d.a.p.r. 0.8.9. n.S.p. 0.p.g. Bd n ae =) NMNZ Ech 375 5 4.0 0.1 1 33 5.8 5 NMNZ Ech 3412 5 3.0 0.1 4 38* = 1 NMNZ Ech 3412a 5 QD) 0.2 4 34 6.8 5 NMNZ Ech 3412b 7 2.0 0.1 2 41 2.7 5 AM J1974 7 1.8 0.1 5 34 3.5 5 AM G7831 7 1.8 0.1 5 48 4.9 4 NMNZ Ech 6 8 1.8 0.1 4 31 4.4 5 BM (NH) 82.12.23.247 (pt) 9 1.3 0.1 5 57* zs 1 AM G11413 9 1.8 0.1 5 ? AM G11032 9 1.5 0.1 5 57* = NMNZ Ech 3413 10 17 0.1 5 72 10.1 5 BM (NH) 82.12.23.247 10 1.6 0.09 5 ? AM J15049 10 1.6 0.09 5 56 3.6 5 NMNZ Ech 3413a 11 1.5 0.1 5 72 10.2 5 AM J15049 11 1.5 0.08 5 56 3.6 5 Abbreviations: d.d., disc diameter; d.a.p.r., dorsal arm plate ratio; 0.s.r., oral shield ratio; n.s.p., number sup- plementary oral plates (maximum available is 5); o.p.g., oral plate granules with mean (x), standard deviation (sd) and number (n) indicated; *, number of granules was estimated due to some missing granules; ?, all oral plate granules were missing (presumably either worn away or dislodged after preservation). Dorsal arm plate ratio was calculated for the fifth dorsal arm plate along the arm. This ratio is an approximate measure of the relative width of that plate’s distalmost margin to its proximalmost one. Oral shield ratio is the approximate length of an individual’s oral shield plate relative to its disc diameter. Bold registration numbers indicate type specimens. After comparing the lectotype of Ophiopeza cylindrica, the lectotype of Ophiopeza gracilis, the holotype and paratype of Ophiopeza arenosa and 11 other specimens from New Zealand and Australia, we conclude that they are conspecific and that their variation in morphology is primarily due to ontogeny. Ophiopeza arenosa and gracilis are consequently synonymized with Ophiopeza cylindrica. O. cylindrica has only been recorded from temperate seas whereas O. spinosa, O. fallax fallax, and O. fallax arabica are only known from tropical seas. In addition to their appar- ent geographic separation, cylindrica and spinosa differ most obviously by the fine, flat- topped granules of spinosa (see p. 275). O. cylindrica differs from fallax fallax and fallax arabica not only by the finer granulation of cylindrica but both fallax and arabica possess shorter arm spines, some bare marginal plates and, 1n arabica, bare radial shields. Ophiopeza spinosa (Ljungman, 1867) Fig. 3A,B Ophiarachna spinosum Lungman, 1867: 305. Ophiopeza fallax. Litken, 1869: 35. [non fallax Peters, 1851.] Pectinura spinosa. Lyman, 1874: 221. Ophiopezella spinosa. Ljungman, 1872: 639. Lyman, 1882: 17. H. L. Clark, 1909a: 120; 1915: 304; 1921: 141, 1946: 258. Koehler, 1922: 338. Marsh and Marshall, 1983: 678. Ophiopezella lutkent de Loriol, 1893a: 392, pl. 13, fig. 1-le. Ophiopezella dubiosa de Loriol, 1893b: 7, pl. 23, fig. 2-2f. H. L. Clark, 1909a: 120. Koehler, 1922: 120. Ophiopeza dubiosa. A. M. Clark, 1968: 313. Clark and Rowe, 1971: 127. Ophiopeza spinosa. A. M. Clark, 1968: 313. Clark and Rowe, 1971: 90, 127, fig. 44e. Gibbs et al., 1976: 130. Kingston, 1980: 145. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 274 OPHIOPEZA AND OPHIOPSAMMUS Material examined: (All specimens examined are held in the AM) 3 spec., J5352, Norwest Id, Capricorn Group, Qld, no coll. depth, vu.29; 3 spec., J5967, Norwest Id, Capricorn Group, Qld, no coll. depth or date; 2 spec., J8881, NW of Gillett Cay, Swain Reefs, Qld, no coll. depth, 17.x.62; 2 spec., J8882, off Gillett Cay Swain Reefs, Old, 65- 7Aimp!3s2625-2)spec:, 9303, Heron 1d) Old) noycoll> depth, x11-592 specs 10959» Marion Reef, Old, 4m, 1.1x.77; 1 spec., J10940, Marion Reef, Old, 8m, 27.vii1.77; 1 spec., J15134, South Solitary Id, N.S.W., 27m, 31.1.82; 1 spec., J16602, Malabar, Lord Howe Id, 10m, 10.xi1.79; 1 spec., J16856, Three Isles, Old, 3-12m, 6.x.82; 3 spec., J19461, Heron Id, Old, no coll. depth, 29.viii.85; 1 spec., J19577, Heron Id, Qld, no coll. depth, 23 Villeoo; specs, 19578, Heron Id} Old) no coll) depth, 25.vin85;slspecsm loans: Heron Id, Old, no coll. depth, 28.viu.85. Fig. 3. A, B, Ophiopeza spinosa, (AM J19461), dorsal and ventral view respectively, dd = 6mm. Some disc granules were removed in A to show disc scaling. Diagnosis: Disc diameter to 9mm, densely covered with small granules which are tri- angular in cross-section and flat-topped with a slightly depressed central area, ca 50 per mm; coarse overlapping scales underlying the granules. Each interradius with about nine marginal disc plates which are convex, prominent, and covered with larger gran- ules than are present on the rest of the disc. Radial shields small (0.1 x d.r.), obscured by granules; only 1-2 scales separating the two plates of each pair of radial shields. Arms to 30mm, ca 5 x d.d., dorsal plates fan shaped, ca as wide as long. Nine to twelve, pointed, arm spines, less than half an arm segment long. Iwo tentacle scales per pore, inner one longest, outer one covering base of lowest arm spine. Oral shields longer than wide with a blunt proximal margin, supplementary plates usually present; adoral plates triangu- lar, ca half as long as the oral shields, not contiguous. Oral plates covered with flat- topped granules. Eight to ten oral papillae; tooth papillae generally pointed. Colour (dried): Dorsal surface cream to light grey, arms generally with dark banding; ventral surface cream, little sign of banding. Colour (live): Kingston (1980: 145) described one live specimen as ‘cream with faint brown cross-bands on the dorsal arm surface’. Distribution: In Australia, from off Coffs Harbour, N.S.W., along the Great Barrier Reef, Old, and in off-shore reefs of northwestern Australia. Overseas, western Indian PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 L. L. VAILANDF. W. E. ROWE P15) Ocean and the Red Sea to Indonesia, the Philippines and the western Pacific Ocean. A cryptic species often found under rocks or rubble, 3-74m. Remarks: Both H. L. Clark (1909a) and Clark and Rowe (1971) were hesitant in recog- nizing spinosa and dubiosa as distinct species. De Loriol (1893b) considered dubiosa separate from spinosa because it had fewer arm spines (9 compared with 12-14) and because of its yellowish-green colour, instead of brown as in spinosa. We have examined 20 specimens of spinosa held in collections of the Australian Museum. Their disc diameters ranged from 4-9mm and their arm spine number from 9-12, overlapping the numbers supposedly characteristic of these two species. Although specimens examined by us were coloured a shade of brown, we do not consider the slight variation in colour recorded by de Loriol to be significant. In addition, our material also agrees with de Loriol’s figures of dubiosa. Thus, we herein consider dubiosa to be synonymous with spinosa. O. spinosa shares with cylindrica, fallax, and arabica the character of only a few large scales separating the two plates of each pair of radial shields. O. spinosa can be separated from these taxa in possessing only 1-2 scales in that position, but more readily by the distinctive shape of its disc granules. Key to the species of Ophiopeza (excluding O. exzlzs) Discrconpletelyacoveredswathy Graril CS sear tes ese eee eye el eee 2 ? Some marginal disc plates, both radial and interradial, without granules ....... 3 Marginal interradial disc plates convex, conspicuous despite cover of granulation; liseqerramulesslAat—tO pp eC. c scien sss ten ey cartes she ces oie Munir ens ep ate O. spinosa 2? Marginal interradial disc plates not conspicuously convex, not visible through Cranuilationacdiscsranules roundedess sae ess: hye Shi ee O. cylindrica 3. Only the middle marginal interradial disc plates are bare; radial shields covered WuailamenccrMUnl syle es suse Sp ater, as MA 12s ene NAT eteveatecl at Seal O. fallax fallax (Fig. 4A, B) 3’ Some marginal disc plates, both radial and interradial, are bare; radial shields Sometinaesppantiallysbaney scm eho sciiscicmenrmueuedanen Scene a. luaen, O. fallax arabica Genus Ophiopsammus Litken Ophiopsammus Lutken, 1869: 37 (type-species Ophiopeza yoldi, Lutken, 1856, by monotypy). Diagnosis: A genus of ophiodermatid with radial shields obscured by granules; margi- nal interradial disc scales enlarged, covered by granules, never bare; disc scales very fine, overlapping, numerous disc scales separating the two plates of each pair of radial shields; oral plates covered by granules; oral shields naked, supplementary plates absent or present; dorsal arm plates transversely rectangular (up to ca 5 x wider than long), convex to conspicuously carinate; arm spines seldom exceeding one arm segment in length, often appressed. Other species included: O. aequalis (Lyman, 1880), O. anchista (H. L. Clark, 1911), O. angusta sp. nov., O. assimilis (Bell, 1888), O. maculata (Verrill, 1869). Remarks: We support A. M. Clark’s (1968) resurrection of Ophiopsammus, but not solely on the basis of its carinate dorsal arm plate shape. In addition, our studies have shown the importance of disc scaling in recognizing Ophiopsammus. This genus differs from PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 276 OPHIOPEZA AND OPHIOPSAMMUS Fig. 4. A, B, Ophiopeza fallax fallax, (BM (NH) 1965.6.1.507), dorsal view, dd = 14mm. Some disc granules were removed to show disc scaling. Ophiopeza in having relatively finer disc scaling, and numerous small scales separating the two plates of each pair of radial shields. In addition, the dorsal arm plates are wide and transversely rectangular shaped. The species we include in Ophiopsammus also attain PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 ) ) L. L. VAILANDF. W. E. ROWE DHT a greater size (d.d. to ca 41mm, arm length to ca 180mm) than those of Ophiopeza (d.d to ca 11mm, arm length to ca 40mm). Ophiopsammus yoldi Lutken, 1856 Fig. 5A,B Ophiopeza yoldu Lutken, 1856: 9. Lyman, 1874: 221. Ophiopsammus yoldi. Lutken, 1869: 37. A. M. Clark, 1968: 317, fig. 9a, b. Clark and Rowe, 1971: 90, 127, pl. 21, figs 7, 8. Gibbs et al., 1976: 130. Kingston, 1980: 145. Ophiopeza conjungens. Bell, 1884: 137. Doderlein, 1896: 281, pl. 15, fig. 1. ecunuawoldim leis Clarkal909a: 191921 4 19383445 Koehlen192237333, 1930: DIO Fig. 5. Ophiopsammus yoldi, (AM J5374), dd = 22mm. A, dorsal view, some disc granules were removed; B, arm spines (proximal) and a portion of dorsal arm plate (top of photo). Material Examined: (All specimens examined are held in the AM) 3 spec., G4208, Mapoon, Gulf of Carpentaria, Old, no coll. depth or date; 1 spec., G4983, Port Curtis, Qld, no coll. depth or date; 1 spec., G11420, Port Denison, Qld, no coll. depth or date; 4 spec., J2413-16, Port Curtis, Old, no coll. depth or date; 2 spec., J5307, Albany Passage, Olde? ime 1x23 S2uspecs oo 7/4) olf Gatcombe Head, Port ‘Gurtis, Old 17-22m, vil.29; 3 spec., J9375, off Gatcombe Head, Port Curtis, Old, 17-22m, vi.29; 1 spec., J5487, Port Curtis, Old, no coll. depth or date; 1 spec., J5924, Kennedy Sound, nr Whitsunday Passage, Old, 15m, no coll. date; 1 spec., J8889, off Gillett Cay, Swain Reefs, Old, 65-74m, 19.x.62; 1 spec., J8890, off Gillett Cay, Swain Reefs, Old, 65-74m, 138x162; 1 spec!, J9306, Gulf of Carpentaria, Old; <26m, 1x.64; 1 ‘spec:, Jll227, SE corner of Gulf of Carpentaria, Qld, no coll. depth or date; 1 spec., J12208, nr Turtle Head Id, Cape York, Qld, no coll. depth, 15.11.79; 12 spec., J12228, Cape York, Old, no coll. depth, 15.1.79; 1 spec., J18001, Abbot Pt, Bowen Qld, 15-17m, 11.v1.83; 5 spec., J18002, Abbot Pt, Bowen, Qld, 6-11m, 111.83; 4 spec., J18009, Abbot Pt, Bowen, Old, 11-12m, 111.83; 2 spec., J18010, Abbot Pt, Bowen, Qld, 15-17m, 11.vi.83; 1 spec., J18013, Abbot Pt, Bowen, Qld, 16m, 12.v1.83; 2 spec., J18016, Abbot Pt, Bowen, Old, 10-16m, 12.vi.83; 2 spec., J18017, Abbot Pt, Bowen, Old, 5-19m, 10.vi.83; 1 spec., J18018, Abbot Pt, Bowen, Qld, 5-19m, 10.vi.83; 2 spec., J18048, Abbot Pt, Bowen, Qld, 5-6m, 19.vi.82; 2 spec., J18049, Abbot Pt, Bowen, Qld, 14-15m, 111.83; 1 spec., J18345, 74 n.m. NNE PROG. LINN. SOG. N.S.W., 110 (3), (1988) 1989 278 OPHIOPEZA AND OPHIOPSAMMUS Pt Hedland, W.A., 80m, 30.x.83; 1 spec., J18370, 50 n.m. NNE Pt Hedland, W.A., 38- 40m, 25.x.83; 1 spec., J18400, 52 nm. NNE Pt Hedland, W.A., 36-37m, 24.x.83; 1 spec., J18409, 52 n.m. NNE Pt Hedland, W.A., 36-37m, 24.x.83; 1 spec., J18478, 80 n.m. NNE Pt Hedland, W.A., 82m, 23.x.83; 1 spec., J18497, 80 n.m. NNE Pt Hedland, W.A., 83m, 23.x.83. Diagnosis: Disc diameter to 22mm, densely covered with rounded granules, 14-23 per mm; fine, overlapping scales underlying the granules. Each interradius with about 9 marginal disc plates, obscured by granule cover, middle plate the largest. Radial shields 0.3 x d.v., obscured by granules, the two plates of each pair of radial shields separated by numerous scales. Arms to 75mm, 1.8-3.1. x d.d.; dorsal arm plates mainly transversely rectangular in shape (2-5 x wider than long) but becoming fan shaped near the arm tip, carinate, even in small specimens, usually entire but occasionally fragmented; ventral arm plates ca as long as wide in small specimens (d.d. = 10mm), becoming wider than long (up to 2 times) in larger specimens, distalmost margin slightly irregular. Five to ten stout arm spines, middle ones longest (sometimes slightly exceeding an arm segment in length) and in large specimens (d.d. 2 20mm) also wider than the remaining spines. Two tentacle scales per pore, inner one larger, ca half the length of lowest arm spine and distinctly curved, outer one covering base of lowest arm spine. Supplementary oral shields generally absent, rudimentary when present. Oral shields pentagonal in small specimens, becoming heart-shaped with growth, flat, ca as wide as long; adoral plates small, triangular, not contiguous. Oral plates covered with granules. Ten to twelve oral papillae; tooth papillae pointed. Colour (dried): Dorsal surface light grey or brown to a darker reddish brown, ventral surface generally a lighter shade of the dorsal surface colour. Some specimens have mottled cream and grey discs with grey banding on cream coloured arms, while others have a large, light stellate pattern in the disc centre contrasting with a darker background. Distribution: In Australia, along the Great Barrier Reef and in the Gulf of Carpen- taria, Old, and off Pt Hedland, W.A.; 5-83m. Overseas, the species has been recorded from the Bay of Bengal to the Philippines Ids, to 215m. Remarks: Housed in the Australian Museum is a collection of approximately 60 speci- mens of Ophiopsammus yoldi and 40 specimens of Ophiopsammus assimilis. Examination of dorsal disc scaling in these specimens has shown the two species to be very similar. In fact, the very distinctive arm spine arrangement of yoldi is the only character we can find to separate it from assimilis. In yoldi, the middle arm spines are longest (ca one arm segment long), whereas in assimilis all arm spines are of approximately equal length (ca 4%-*%, of an arm segment long). Characters separating yoldu from its congeners are given in the key. Ophiopsammus aequalis (Lyman, 1880) Fig. 6A-C Ophiopeza aequalis Lyman, 1880: 9, pl. 2, figs 23-25; 1882: 12, pl. 27, figs 7-9. Koehler, 1904: 10; 1922: 337, pl. 77, figs 16, 17. A. M. Clark, 1968: 313. Pectinura aequalis. H. L. Clark, 1909a: 118; 1915: 303. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAIL AND F. W. E. ROWE 279 Fig. 6. Ophiopsammus aequalis, (AM J13960; 2 specimens, 1 and 11). A, dorsal view, specimen 1, dd = 23mm; B, dorsal view, specimen 11, dd = 26mm; C, arm spines (proximal) and ventral arm plates (bottom of photo), specimen 1. Some disc granules were removed in A and B. Material examined: 3 spec., AM J13960, Timor Sea, northern Australia (09° 48°S, L222" 13), LIV AT aay, M5 )sae 7S) Diagnosis: Disc diameter to 23mm, densely covered with small, round granules, ca 15 per mm; exceedingly fine overlapping scales underlying the granules. Each interradius with eleven marginal interradial disc plates, slightly convex, middle one largest, all PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 280 OPHIOPEZA AND OPHIOPSAMM US covered by granules. Radial shields 0.15-0.2 x d.r., obscured by granules; numerous scales separating the two plates of each pair of radial shields. Arms to ca 140mm (tips broken), ca 5-6 x d.d.; dorsal arm plates transversely rectangular (2-3 x wider than long) and strongly carinate. Nine to ten pointed arm spines, %-% of an arm segment long, lowest spine marginally longer and wider than the remaining spines. Two tentacle scales per pore, inner one longer, outer one covering base of lowest arm spine. Oral shields slightly wider than long, proximal margin blunt, distal margin broadly convex, some oral shields with a rudimentary supplementary plate; adoral plates small, ca 0.2-0.3 x oral shields long, not contiguous. Oral plates covered by numerous granules. Eight to nine oral papillae, penultimate very long (up to 3 x longer than the other papillae); tooth papillae blunt. Colour (dried): Cream to light grey both dorsally and ventrally. Koehler (1922: 337) records one preserved specimen with a pink dorsal disc surface and arms with light pink and dark red annulations. Distribution: Lyman’s types came from northeast of New Guinea and the Kei Islands. Additional specimens were collected by the ‘Siboga’ northeast of northern Borneo. The three specimens examined by us from the Timor Sea are the first records of this species from Australia. This is a deep-water ophiodermatid species previously collected from 209-274m. Remarks: Long delicate arms are common to both O. aequalis and O. angusta sp. nov., in contrast to the relatively shorter, stouter arms of O. assimilis and O. yoldi. However, aequalis is easily separated from angusta on differences in arm spine arrangement and from the New Zealand species O. maculata on the shape of dorsal arm plates. Characters separating aequalis from its congeners are illustrated in the key. Ophiopsammus assimilts (Bell, 1888) Fig. 7A-C Pectinura assimilis. Bell, 1888: 282, pl. 16, fig. 5. H. L. Clark, 1909a: 118; 1946: 257. A. M. Clark, 1966: 327. ?Baker, 1982: 436. ? Ophiopeza cylindrica. Farquhar, 1898a: 190; pl. 14, figs 4, 5 [non cylindrica Hutton, 1872.] Pectinura dyscrita. H. L. Clark 1909b: 534, pl. 49, figs 5-7. Dartnall, 1980: 42, 71. H. L. Clark, 1946: 256. ? Pectinura cylindnca. Mortensen, 1924: 172, fig. 35(1-2) [non cylindrica Hutton, 1872.] Fectinura nigra. H. L. Clark, 1938: 344. H. L. Clark, 1946: 256. Ophiopeza dyscrita. A. M. Clark, 1968: 313. Baker, 1982: 436. Ophiopeza nigra. A. M. Clark, 1968: 313. Material examined: Holotype of assimilis BM(NH) 86.6.9.23, Port Jackson, N.S.W., no coll. depth or date; holotype of nigra MCZ 5257, Koombana Bay, Bunbury, W.A., 9- 15m, 26.x.1929; 1 lectotype of dyscrita AM J849, off Wata Mooli, N.S.W., 129-144m, no coll. date; 2 spec., AM E5941, 30 mls S. of Mt Cann, Gippsland, Vic., 129-185m, 19.x.14; 1 spec., AM J11637, Long Reef, Collaroy, N.S.W., 40m, 14.1v.72; 1 spec., AM G11418, Port Jackson, N.S.W., no coll. depth or date; 4 spec., AM J15991, (32° 52.51°S, 152° 34.35’E), 151m, 6.xii.78; 1 spec., AM G11440, Port Jackson, N.S.W., no coll. depth or date; 1 spec., AM Ji0008, E of Cronulla, N.SW. (34° 09’S, 151° 16’E), 127-132m, 17.1v.75; 3 spec., AM J14978, Julian Rocks, Byron Bay, N.S.W., 12m, 4.11.82; 1 spec., AM J18557, nr Mistaken Id, Albany, W.A. (35° 04’S, 117° 56’E), 2m, 13.x.83; 1 spec., PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAILAND FE. W. E. ROWE 281 Fig. 7. Ophiopsammus assimilis, holotype (BM (NH) 86.6.9.23), dd = 24mm. A and B, dorsal view, some disc granules were removed: C, ventral arm plates and arm spines (proximal). AM J14979, Julian Rocks, Byron Bay, N.S.W., 9-15m, 3.11.82; 1 spec., AM E4717, Gabo Id, Vic., 368m, no coll. date; 1 spec., AM G7115, Port Hacking, N.S.W., 55-74m, 1961; 1 spec., AM J12564, NE side Kangaroo Id, S.A., 5m, 9.11.78; 1 spec., AM G11419, Port Jackson, N.S.W., no coll. depth or date; 1 spec., AM J17621, Triggs Id, Perth, W.A., 6m, 221.83; 3 spec., AM E5212, E of Babel Id, Tas., 129m, 1914; 4 spec., AM J11086, 40mls ENE of Sydney, N.S.W., 184-232m, 18.v1.59; 1 spec., AM G11027, Port Jackson, N.S.W., no coll. depth or date; 2 spec., AM J13975, Montague Id, N.S.W., 28-31m, 16.11.81; 1 spec., AM J14159, Ulladulla, N.S.W., 25m, 27.11.81; 1 spec., AM J13991, Montague Id, N.S.W., 31m, 16.11.81; 1 spec., AM J14015, Burrewarra Pt, N.S.W. (35° 50’S, 150° 14’E), 28m, 15.11.81; 2 spec., AM J14162, Jervis Bay, N.S.W., 25-28m, 21.111.81; 3 spec., AM G2032, Wellington, N.Z., no coll. depth, x1.1896; 2 spec., AM G11423, Port PROC. LINN. SOC. N.S.W,, 110 (3), (1988) 1989 282 OPHLOPEZA AND OPHIOPSAMMUS Jackson, N.S.W., no coll. depth or date; 1 spec. AM J4306, Shell Harbour, N.S.W., no coll. depth, 21.11.77. Diagnosis: Disc diameter to 28mm, densely covered with rounded granules, 8-15 per mm, very fine, overlapping scales underlying the granules. Each interradius with 8-14 marginal plates, all concealed by granulation. Radial shields 0.2 x d.r., obscured by granules, up to seven overlapping scales separating the two plates of each pair of radial shields. Arms to 100mm, 2.3-4.7 x d.d.; dorsal arm plates convex to carinate, trans- versely rectangular (1.7-2.7 times wider than long); ventral arm plates as wide as long. Arm spines stout, tapering, slightly swollen at the base, ca half an arm segment long, 4- 11 spines per lateral plate. Two tentacle scales per pore, inner one the longer. Oral shields usually without a supplementary oral plate, poorly developed if present; adoral plates small, not contiguous. Oral plates covered with granules. Seven to eight blunt oral papillae on each half jaw; penultimate conspicuously wider than the others; tooth papillae stout, blunt. Colour (dried): Variable, cream to reddish brown to grey; arms sometimes with faint banding. Colour (live): Some specimens may have a bright red disc and arms with alternating bands of red and grey while others have a red/cream mottled disc. Distribution: A few individuals have been collected near Perth and Albany, W.A. and one from Kangaroo Id, S.A. This species has been more commonly found in ‘Tas. and from off Gabo Island, Vic., to Byron Bay, N.S.W., on the Australian coast. It also occurs in New Zealand, at least along the northern coasts from Three Kings Id to Cook Strait. This species is quick moving and is found under rocks (pers. obs. L.V.), at least in shallower depths; depth range 2-368m. Remarks: The original description of Ophiopeza assimilis Bell, 1888, is based on one specimen with a disc diameter of ca 100mm and arm length of 24mm. We have examined a specimen from the British Museum (86.6.9.23) labelled as a type of asszmilis with a disc diameter of 24mm and arm length of ca 90mm. We consider this specimen the holotype of asszmzlis since it fits Bell’s description and has the same locality data. Only its disc diameter and arm length measurements disagree with the original descrip- tion and these have obviously only been reversed. As discussed previously (p. 272), figures given for Ophiopeza cylindrica by both Farquhar (1898a) and Mortensen (1924) probably represent assimilis. This view 1s sup- ported by material in the Australian Museum collections from New Zealand (AM G2032). Positive identification of the material figured by them would have required removal of some dorsal disc granules in order to reveal the pattern of disc scaling between the plates of each pair of radial shields, but this material was not available. In a revision of the genus Pectinura, H. L. Clark (1909a) separated arenosa (now regarded as a synonym of Ophiopeza cylindrica) and assimilis on the size of their dorsal arm plates and on the presence or absence of supplementary oral shields. Specimens described herein as cylindrica have supplementary oral plates associated with most oral shields (Table 1). These plates are generally absent in assimilis, although some in- dividuals (AM; G7115, J14978 (2), E5941) have rudimentary supplementary oral plates. The dise granules of cylindrica and assimilis have never featured in descriptions of these species and consequently two characters which readily separate them have been over- PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAILAND F. W. E. ROWE 283 looked: namely the arrangement of scales separating the two plates of each pair of radial shields, and the type of disc scaling. Shortly after his revision of Pectinura, H. L. Clark (1909b) described a new species, dyscrita, from two specimens collected off central N.S.W. (lectotype AM J5849; para- lectotype MCZ 590, designated herein). He noted they were most similar to asszmilis but differed in having fewer (6-8) arm spines. Another species, nzgra, was described by H. L. Clark (1938) from one specimen collected near Bunbury, Western Australia. H. L. Clark (1946) separated nigra and dyscrita on arm spine number and by the black coloration of nigra. We have examined the holotype of nigra (d.d. 7mm) and the lectotype of dyscrita (d.d. 10mm) and confirm that their disc scaling is the same as that of asszmalzs. Further- more, differences in their arm spine number and coloration are within the range of vari- ation found in assimilis. However, as more material is collected, it may be shown that the darker coloration which Clark attributed to nigra is more common in specimens from South Australia and westwards. H. L. Clark (1909; 1946) also distinguished dyscrita from other species of Ophiopeza by its large lowest arm spine. Our examination of the lectotype material of dyscrita has shown its lowest arm spine is only marginally longer, if any, than the other arm spines. Thus, we now regard both dyscrita and nigra as synonyms of assimilts. Interestingly, an enlarged lowest arm spine is one character readily separating adults of Ophiopsammus angusta sp. nov. (p. 285) and assimilis. In addition, the lowest arm spine of angusta is broader than the other spines and its arm spines have straight margi- nal edges in contrast to the tapering ones of asszmilis. O. assimilis is most readily separated from its other congeners on arm spine arrangement and dorsal arm plate shape as shown in the key. Large New Zealand specimens, however, occasionally have frag- mented dorsal arm plates (A. N. Baker, pers. comm.). Ophiopsammus angusta sp. nov. Fig. 8A-C Material Examined: Holotype, AM E4693, E of Flinders Id, Bass Strait, ‘Tas., 148- 553m; no coll. date; 1 paratype, AM E5025, E of Maria Id, Tas., 144m, no coll. date; 1 paratype, Am E5376, NE of Cape Pillar, Tas., 148m, 1914; 1 paratype AM E5946, Bay of Fires, Tas., 102m, 7.xi.14; 1 paratype, MV F52197, S of Cape Nelson, Vic., 406m, 10.11.77; 1 spec., AM E5116, E of Maria Id, Tas., 236m, no coll. date; 10 spec., MV Boz OMBassiStraity 36-9220) 5; 1482 29:2" EF) 140m, 1o:x1.81; I spec. MV F521947 Sof BemumeRaver Vicw o>) lone) 149°%00" E223, 22x03; 1 specs NIV" Bo219 5s Wok Cape Nelson, Vic., 165-201m, 6.vi.69. Diagnosis: Disc diameter to 2imm; rounded granules, 7-14 per mm; fine overlapping scales on the dorsal and ventral surfaces; 9-14 marginal disc plates per interradius, middle one largest. Radial shields ca 0.1-0.2 x d.r., about twice as long as wide, the two plates of each pair separated by ca 8 disc scales, radial shields obscured by disc granu- lation. Arms to 116mm, 5.5-7.9 x d.d., dorsal arm plates transversely rectangular (ca 1.5 x wider than long), slightly carinate, distal margins slightly convex; ventral arm plates roughly hexagonal with rounded corners (ca as wide as long), distal margin convex; 8-10 arm spines, ca half the length of an arm segment; lowest arm spine the longest, conspicu- ously wider than the other arm spines. Lateral margins of arm spines generally parallel, tapering on the distal 4% of their length. Two tentacle scales per pore, inner one the longer. Oral shields roughly triangular with round edges, supplementary plates usually absent; adoral plates longer than wide, not contiguous. Oral plates covered with PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 284 OPHIOPEZA AND OPHIOPSAMMUS Fig. 8. A-C, Ophiopsammus angusta sp. nov., holotype (AM E4693), dd = 17.5mm. A and B, dorsal view, some disc granules were removed; C, ventral arm plates and arm spines (proximal). granules. Six to seven oral papillae on each half jaw, penultimate at least 2 x wider than the remainder; tooth papillae stout, blunt. Description of Holotype: Disc diameter 17.5mm; covered with granules both dorsally and in the ventral interradial areas, 7-9 granules/mm. Fine, overlapping disc scales occur both on the dorsal and ventral surfaces. Each interradius with thirteen marginal disc plates, middle one the largest. Radial shields 0.2 x d.r. (ca twice as long as wide), the two plates of each pair separated by ca 8 disc scales. Disc scales, marginal plates and radial shields obscured by disc granulation. Arms broken, to ca 100mm, 5.7 x d.d., very finely tapered, slightly carinate dorsally. Dorsal arm plates transversely rectangular with a slightly convex distal margin (ca 1.5- PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAILAND F. W. E. ROWE 285 1.7 x wider than long), distally becoming more fan shaped with a convex distal margin. First ventral arm plate transverse diamond shaped (ca 2 x wider than long), distal margin convex. Succeeding arm plates roughly hexagonal with rounded corners (ca as wide as long), distal margin convex; distal plates becoming slightly longer than wide. First lateral arm plate with 2 arm spines, 2nd and 3rd-4, 4th and 5th-6, 6th-8, thereafter 9, decreasing at the distal end of the arm to 6 spines. Lowest arm spine longest (34-1 arm segment long) and widest (ca 1.25 x wider than the other spines). Remaining arm spines ca % an arm segment long, proximal % of each spine with parallel lateral margins, becoming slightly tapered distally with a blunt tip; adjacent spines contiguous or almost contiguous. ‘Iwo tentacle scales per pore; inner one the longer, tapering distally; outer one covering proximal portion of lowest arm spine (ca as broad as long), with a blunt distal margin. Oral shields roughly triangular with very rounded edges (ca as long as broad), one shield with a rudimentary, distal, supplementary plate; adoral plates triangular (longer than wide); not contiguous. Oral plates densely covered with granules about equal in size to the disc granules. Six to seven oral papillae on each half jaw, penultimate ca 3 x wider than the other papillae. ‘Tooth papillae stout with pointed or blunt tips. Paratypes: Disc diameters ranged from 11-21mm and maximum arm lengths from ca 60-116mm. Except for overall size and arm spine number, the paratypes do not differ from the holotype. Arm spine number ranged from 8-10 per plate, with the smallest specimen (d.d. 11mm) having the fewest arm spines and the largest (d.d. 21mm) the most. Colour (dried): Holotype, creamy white; other material, creamy white to light grey; one specimen (NMV F52195) is a uniform light purple. Distribution: Endemic to southeastern Australia; off Tas. and Vic.; 102-201m. Etymology: The species is named from the Latin angustza for the straight sided arm spines. Remarks: O. angusta appears most closely related to asszmilis. Characters separating them are: the R/d.d. ratio; shape of the lowest arm spine; and shape of the other arm spines. In angusta, R/d.d. is 5.5-7.9 for individuals with a d.d. >9mm while in similar- sized specimens of asszmilis, this ratio is 2.3-4.7. Consequently, the arms of angusta appear delicate while those of assimilis seem stout and robust. The lowest arm spine of angusta 1s noticeably longer, wider, and more robust in appearance than other arm spines whereas in asszmilis, this spine is only slightly, if any, longer and wider than the other spines. In larger specimens of angusta (d.d. >9mm), lateral margins of arm spines are nearly parallel, except near the tip where they begin to taper. In contrast, specimens of assimilis (d.d. > 9mm) have more tapered arm spines. In addition, lateral margins of arm spines in angusta are almost contiguous whereas in assimilis they are separated. We have been unable to satisfactorily separate small specimens (d.d. < 9mm) of angusta and assimilis as their R/d.d. ratio is very similar (ca 3.0-4.0) and their arm spines appear similar. Relative size of the lowest arm spine is most useful when separating small specimens. A collection of 9 small specimens (d.d. 6-8mm) of Ophiopsammus from Bass Strait (MV F52196) are probably angusta because of their long and wide innermost arm spines. O. angusta shares with aequalis and anchista a high R/d.d. ratio and thus delicate- appearing arms, although the very carinate dorsal arm plates in aequalis readily separate PROC. LINN. SOG. N.S.W., 110 (3), (1988) 1989 286 OPHIOPEZA AND OPHIOPSAMMUS Fig. 9. A, Ophiopsammus anchista, holotype (USNM 25645), dorsal view, dd = 13mm. B, Ophiopsammus maculata, (AM J10208), dorsal view, scale ine = 10mm. it from these two other taxa. Differences between angusta and its other congeners are demonstrated in the key. Notes on Ophiopsammus anchista (H. L. Clark, 1911) During this study, we examined a holotype (USNM 25645, Fig. 9A) and two para- types (USNM 26214) of anchista H. L. Clark, 1911 from Japan. The species is referable to Ophiopsammus. However, the smallest paratype (d.d. = 4mm) is misidentified, being identified by us as a species of the genus Cryptopelta H. L. Clark, 1909a. O. anchista is most readily separated from its other congeners on the shape of its arms, dorsal arm plates, and arm spines, as shown in the key. Key to the species of Ophiopsammus 1 Dorsal arm plates often very fragmented, usually spotted. ..... O. maculata (Fig. 9) 1’ Dorsal arm plates entire, except for O. yoldit which may have a few fragmented plates; motispotteder nar) sj. /ejsr aa snide sn a vies ae eee a ee 2 Jeeacins lone. delicate appearance (IX/didtratio.o-o 1neadullts) vere eee ee 3 J vans Lelatively short istoutappearance (R/didy ration. 8-45/))) ice aan eee 5 See Dorsal auimaplatesssiionehyacaninates we wen kina eo en ee ease O. aequalis ne Dorsalarmyplatesroundtojonlyslichthycanimates, -p erat oe ee 4: 4 Lowest arm spines distinctly longer and wider than adjacent arm spines; lateral margins of arm spines parallel on proximal % of spine; arm segment no. 10 with CayO-ANMUS PALES acheet ens a8 2 Aaa ios aes teak ge im ow ey ae a CHEMO Tae O. angusta 4 lowest arm spines ca the same length as adjacent arm spines; arm spines tapered, ALYSE SIME nt 10 m0 wibhicalOsa Gunns PING Sweetie cys ic ice ieae O. anchista 5 Arm spines on each segment of ca same length, about % an arm segment long .. O. assimilis 5’ Middle arm spines the longest, to about one arm segment long.......... O. yoldu ACKNOWLEDGEMENTS Miss A. M. Clark, British Museum (Natural History), London, U.K., Prof. R.., Woollacott, Museum of Comparative Zoology, Harvard, Mass., U.S.A., Dr A. N. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 L. L. VAILANDF. W. E. ROWE 287 Baker, National Museum of N.Z., Wellington, N.Z., Dr Suzanne Boyd, National Museum, Melbourne, Victoria, Australia, are thanked for the loan of type specimens. Dr F. W. E. Rowe wishes to acknowledge support for this project from both the Australian Research Grants Scheme (ARGS D18015325) and Marine Sciences and Technologies Grant Scheme (MST 84/2092). We wish to thank Ms Kate Lowe, Photo- graphic Department, Australian Museum, and Mr Alan Howard, Northern ‘Territory Museum, for photographing specimens illustrated in this work and Mrs L. Watt, North- ern ‘Territory Museum, for typing the manuscript. Finally, we wish to thank Ms A. Hoggett, N'T. Museum, and Dr A. N. Baker, National Museum of N.Z., for reading and commenting on the manuscript. References BAKER, A. N., 1982. — Brittle-stars (Class Ophiuroidea). Jn. S. A. SHEPHERD and I. M. THOMAS, (eds), Marine Invertebrates of South Australia. Part I: 418-437. Adelaide: South Australian Govt Printer. BELL, J. F., 1884. — Echinodermata. Jn R. W. COPPINGER: Report on the zoological collections made in the Indo- Pacific Ocean during the voyage of H.M.S. Alert} 1881-2: 117-177, 509-512. London: British Museum (Natural History). ——., 1888. — Descriptions of four new species of ophiuroids. Proc. zool. Soc. Lond.: 281-284. CiarK, A. M., 1966. — Port Phillip Survey 1957-1963, Echinodermata. Mem. Nat. Mus. Vict. 27: 289-384. — , 1968. — Notes on some tropical Indo-Pacific Ophiotrichids and Ophiodermatids (Ophiuroidea). Bull. Brit. Mus. (nat. Hist.), Zool. 16(7): 277-322. , and Rowe, F. W. E., 1971. — Monograph of shallow-water Indo —west Pacific echinoderms. London: British Museum (Natural History). CiarK, H. L., 1909a. — Notes.on some Australian and Indo-Pacific echinoderms. Bull. Mus. comp. Zool. Harv. 52(7): 107-137. ——.,, 1909b. — ‘Thetis’ echinoderms. Mem. Aust. Mus. 4: 519-564. ——, 1911. — North Pacific ophiurans in the collection of the United States National Museum. Bull. U.S. natn. mus. 75: v-xvi, 1-302. ——., 1915. — Catalogue of recent ophiurans: Based on the collection of the Museum of Comparative Zool- ogy. Mem. Mus. comp. Zool. Harv. 25(4): 165-376. ——, 1921. — The echinoderm fauna of Torres Strait: its composition and its origin. Pap. Dep. mar. Biol. Carnegie Instn Wash. no. 10: 1-223. ——.,, 1938. — Echinoderms from Australia. Mem. Mus. comp. Zool. Harv. 55. ——.,, 1946. — The echinoderm fauna of Australia. Washington, D.C.: Publs Carnegie Instn no. 566: 1-567. DARTNALL, A., 1980. — Tasmanian Echinoderms. Univ. of Tasmania fauna handbook no. 3: 1-82. DODERLEIN, L., 1896. — Bericht tber von Herrn Prof. Semon bei Amboina und Thursday Island gesammt- ten Ophiuroidea. Denkschr. med.-naturw. Ges. Jena 8: 279-300. FARQUHAR, H., 1895. — Notes on some New Zealand echinoderms. Trans Proc. N.Z. Inst. 1894, 27: 194-208. ——.,, 1898a. — A contribution to the history of N.Z. echinoderms. /. Linn. Soc. Zool. 26: 186-196. — ., 1898b. — On the echinoderm fauna of New Zealand. Proc. Linn. Soc. N.S.W. 3: 300-327. FORBES, E., 1842. On the radiata of the eastern Mediterranean. Part I. Trans Linn. Soc. Lond. 19(1): 143-153. G1Bps, P. E., CLARK, A. M., and CLARK, C. M., 1976. — Echinoderms from the northern region of the Great Barrier Reef, Australia. Bull. Brit. Mus. (Nat. Hist.), Zool. 30(4): 103-144. HuTTon, F. W., 1872a. — Catalogue of the Echinodermata of New Zealand. Col. Mus. and geol. Surv. Dept, N.Z.: vi + 17. ——.,, 1872b. — Descriptions of some new star-fishes from New Zealand. Proc. zool. Soc. Lond.: 810-812. KINGSTON, S. C., 1980. — The Swain Reefs Expedition: Ophiuroidea. Rec. Aust. Mus. 33(3): 123-147. KOEHLER, R., 1904. — Ophiures de mer profonde. Szboga’ Exped. 45: 1-176. ——., 1905. — Ophiures littorales. ‘Svboga’ Exped. 45b: 1-142. ——,, 1922. — Ophiurans of the Philippine Seas. Bull. U.S. natn. Mus. 100(5): x + 486. ——., 1930. — Ophiures recueillis par le docteur Th. Mortensen dans les mers d’Australie et dans l’Archipel Malais. Vidensk. Meddr. dansk naturh. Foren. 89: 1-295. LJUNGMAN A. V., 1867. — Ophiuroidea viventia huc usgue cognita enumerat. Ofvers. K. Vetensk.-Akad. Forh. 23(9): 303-336. ——., 1871. — Forteckning ofver uti Vestindien of Dr. A. Goes samt under Korvetten Josefinas expedition 1 atlantiska oceanen samlade Ophiurider. Ofv. Kongl. Vet-AK Forh. 6: 615-657. LORIOL, P. de., 1893a. — Echinodermes de la Baie dAmboine. Revue suisse Zool. 1: 359-426. ——, 1893b. — Catalogue raisonné des echinodermes. III. Ophiurides et astrophytides. Mem. Soc. Phys. et Fist. nat. Geneve 32, 1(3): 1-63. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 288 OPHIOPEZA AND OPHIOPSAMMUS LUTKEN, C. F., 1856. — Bidrag til kundskab om Slangestjernerne. II. Ovesigt over de vestindiske Ophiurer. Vidensk. Meddr. dansk. naturh. Foren 1: 1-19. ——,, 1869. — Additamenta ad Historiam Ophiuridarum. 3. Beskrivende og kritiske Bidrag til kundskab om Slangestjernerne. K. danske Vidensk. Selsk. Skr. 5(8): 24-109. LyMAN, T., 1874. — Ophiuroidea and Astrophytidae, old and new. Bull. Mus. comp. Zool. 3: 221-272. ——., 1879. — ‘Challenger Ophiuroidae and Astrophytidae. pt. 2. Bull Mus. comp. Zool. 5: 17-83. ——., 1880. — A structural feature hitherto unknown among echinodermata found in deep-sea Ophiurans. Anniv. Mem. Boston Soc. nat. Hist.: 1-12. — , 1882. — Report on the Ophiuroidea. ‘Challenger’ scr. Results: Zoology, 5: 1-386. MarsuH, L. M., and MARSHALL, J. I., 1983. — Some aspects of the zoogeography of north-western Aus- tralian echinoderms (other than holothurians). Bull. mar. Sci. 33(3) 671-687. MORTENSEN, T., 1924. — Echinoderms of New Zealand and the Auckland—Campbell Islands. II. Ophiuroidea. Vidensk Meddr. dansk. naturh. Foren. 77: 91-177. ——., 1940. — Echinoderms from the Iranian Gulf. Asteroidea, Ophiuroidea and Echinoidea. Dan. scient. In- vest. Iran, part 2: 55-110. Perers, W., 1851. — Ubersicht der von ihm an der Kuste von Mossambique eingesammelten Ophiuren, unter denen sich zwei neue Gattungen befinden. Ber. K. preuss. Akad. Wiss.: 463-466. Rowe, F. W. E., and VaIL, L. L., 1982. — The distributions of Tasmanian echinoderms in relation to southern Australian biogeographic provinces. Jn: J. M. LAWRENCE, (ed.), Echinoderms: Proceedings of the International Conference, Tampa Bay: 219-225. Rotterdam: A. A. Balkema. PROC. LINN. SOC. N.SW., 110 (3), (1988) 1989 Pycnodithella harveyi, anew Australian Species of the ‘Iridenchthoniidae (Pseudoscorpionida: Arachnida) CLARICE M. A. KENNEDY KENNEDY, C. M. A. Pycnodithella harveyi, a new Australian species of the Triden- chthoniidae (Pseudoscorpionida: Arachnida). Proc. Linn. Soc. N.S.W. 110 (3), (1988) 1989: 289-296. A new species of the genus Pycnodithella Beier, P harveyi is described from the Sydney region. It is only the second known species of the genus and is characterized by its relatively small size and a distinctly reticulated carapace. Clarice M. A. Kennedy, School of Biological Sciences, Macquarie University, North Ryde, Australia 2109; manuscript received 15 March 1988, accepted for publication 20 July 1988. INTRODUCTION The family ‘Tridenchthoniidae Balzan 1891 is at present represented by 16 genera including approximately 70 species. Of these, only two species namely: Anaulacodithella australica Beier and Heterolophus australicus Beier, have been described from Australia, the remainder are mostly confined to SE Asia and Africa. The species described herein is only the second known species of the genus Pycnodithella Beier and its first Australian representative. Measurements are based on the examination of 10 specimens of each stage made in accordance with those discussed in Chamberlin (1931). Those in parentheses are the female and follow those of the male. Abbreviations for cheliceral trichobothria and setal formulae follow those devised by Chamberlin (1931). All specimens are preserved in spirit. SYSTEMATIC DESCRIPTION TRIDENCHTHONIIDAE Balzan 1891 Remarks: Chamberlin and Chamberlin (1945) state that the name Dithidae, which has been used by various authors, is a synonym of the Tridenchthoniidae. 1947 Pycnodithella Beier Type species: Pycnodithella abyssinica (Beier) 1944 Emended diagnosis: Dorsum of chelal hand without transverse furrow; fixed chelal finger with 8 trichobothria, et near ds, distinctly separated from it/st; movable chelal finger with acute teeth extending to finger base; intercoxal tubercle absent. Remarks: The genus Pycnodithella was erected by Beier (1947: 287) solely on the basis of the position of the trichobothria e¢ in relation to the double setae ds. The discovery of a new species, here assigned to Pycnodithella has therefore necessitated the emended diagnosis for the genus. Pycnodithella differs from Verrucadithella Beier 1931 in that the dorsum of the chelal hand does not possess a transverse groove. Anaulacodithella (Beier) 1944 is distinguished from Pycnodzthella in that the trichibothria et, zt and est of the fixed chelal finger are grouped togoether, distinctly separate from the double setae ds and that the teeth of the movable chelal finger, prox- imal to the trichobothria st, are replaced by a wavy lamella extending to the finger base. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 290 NEW AUSTRALIAN PSEUDOSCORPION 10.un® Fie. 1. Scanning electron micrographs. Pycnodithella harveyi n. sp., male paratype: A, lateral aspect of left chela; B. fixed chelal finger showing trichobothria ¢ near ds; C, movable chelal finger showing trichobothria ¢, stand ‘4 equidistant; D, chelicerae, dorsal aspect; E, distal lamellae of serrula exterior; F, Carapace showing conspicuous reticulate sculpture and well developed eyes; G, unilateral pinnate blades of flagellum; H, pretarsus PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 C. M. A. KENNEDY 291 Pycnodithella harveyi sp.nov. (Ene Wy Ene, A, Leer, ah, Whey. Ge) Etymology: This species is named in honour of Dr M. S. Harvey in recognition of the great contribution he has made to the knowledge of the Pseudoscorpion fauna in Australia. Holotype: ©, Australian Museum, Sydney (KS 17441). Type locality: Macquarie University campus, North Ryde, 13.5km NW of Sydney CaP@PINES WE 33247 7S) 1512077 En in litters 17-vie8 5, CK ennedy, Paratypes: same data as holotype, Australian Museum, 60° (KS 17442), 69 (KS 17443), 6 nymphs (KS 17444). Distribution to various other museums in progress. Diagnosis: Irichobothria ¢, st and sb equidistant (Fig. 1C); fixed chelal finger with 20- 22, movable chelal finger with 16-18 acute, well-spaced teeth. Carapace conspicuously reticulate; chela (with pedicel) 0.59-0.64 (0°), 0.64-0.73 (9) mm in length, 4.20-5.08 (0°) 4.0-4.70 (9 ) x longer than broad; 1st and 2nd pedal coxae each with 4 medial coxal spines. Description: Adults. Colour dark brown; pleural membrane cream, smooth; dorsal surface of cephalothorax with conspicuous reticulate sculpture extending to cornea of eyes and tergites (Fig. 1F), fields of reticulation deeply depressed on carapace, some- what shallow on tergites. Pedipalpal trochanter with gently rounded anterior margin, L/W ratio 1.10-1.71 (O), 1.20-1.80 (9 ), femur straight increasing in width distally 3.02- 2.50 (O°), 3.83-4.80 (Q), tibia pyriform, curved towards tibio-femoral insertion, 1.90- 2.30 (0), 1.75-2.25 (Q); chela distally setose, very slightly papillate dorsally, smooth ventrally with 20-30 short, acuminate setae, chela (with small pedicel) 4.21-5.08 (©), 4.05-4.71 (9), chela (without pedicel) 4.0-4.90 (0"), 3.52-4.50 ( Q ) x longer than broad. Fixed finger with 8 trichobothria, ef near ds and distinctly separate from zt/est (Fig. 1B), movable finger with 4 trichobothria (Fig. 1A), marginal teeth acute, distinctly separated on both fingers, progressively smaller proximally, never contiguous, 20-22 fixed finger, 16-18 movable finger (Fig. 3R); 2 minute teeth at base of terminal teeth precede mar- ginal rows. Chelicerae large, stout, fixed finger dorsally papillate except distally, broad smooth band along posterior margin, 6-8 setae, /s absent, 10 minute antrorse teeth, first large; movable finger slender, galeal seta anterior to 7-8 marginal teeth, terminal tooth large, erect, others progressively very small and antrorse, galea absent. (Fig. 1D). Serrula exterior with 16-17 (0”), 15-16 (9 ) lamellae (Fig. 1E); serrula interior with 8-10 thick, blunt lamellae, serrate margin. Flagellum with 8-10 slender, unilaterally pinnate blades of sub-equal length, distally divided into 3-4 short spines that are bent at an obtuse angle with respect to the blades and are directed towards the midline; 6-7 long ‘spines’ well-spaced along the upper two thirds of the blades point in the same direction (Fig. 1G); base of blades stalk-like. Carapace (Fig. 1F) anterior margin serrate, median blunt vestigial epistome (Fig. 3P); setae spine-like 8:8-10:50-60 ( 0”), 8:8-10:40-50 ( 9 ); two well-developed eyes; L/W ratio 1.0-1.16 (0°), 0.77-1.0 (Q) x longer than broad. Tergal chaetotaxy: O 12-14: 12-14: 14-16: 14-16: 15-16: 15-16: 16-18: 15-16: 10-12: 10: 8-10: 2; Q 12-14: 12-14: 12-14: 14-16: 14-16: 14-16: 13-16: 12-15: 13-14: 10-12: 10: 2, uniseriate. Coxae: Ist pedal coxae with anterior mesial process; Ist and 2nd pedal coxae each with 4 short medial coxal spines transversely incised into finger-like projections distally (Fig. 2I, J), base of spine thick; intercoxal tubercle absent. Coxal chaetotaxy: O 2: 4-5: 6-8, OsO= 84", OF OBR Se), OB O=68 PMO), Os tsk teiaIliie O) We Oe D(0y, Oe Geez 40), Wa Zee Ghee eto Gs), 0: 4-6: 10-12, 8-10 long acuminate setae on posterior border. Male genitalia: consisting of complex series of sclerotized apodemes and rods, dorsal anterior apodeme appears brush-like; ejaculatory canal atrium cup-shaped and anterior to genital armature (Fig. 4V). Fig. 4T shows the three dimensional structure of the genitalia. Chaetotaxy: C PROC. LINN. SOG. N.S.W., 110 (3), (1988) 1989 292 NEW AUSTRALIAN PSEUDOSCOR PION Fig. 2. Pycnodithella harveyi n. sp. 1, male-coxal spines; J, enlargement of distal portion of one coxal apine; K, male genital region, external aspect; L, female genital region, external aspect; M, tritonymph showing multiple simple galeae. genital plate medially in the form of a diamond; anterior operculum with 5 marginal, long acuminate setae each side projecting into open cavity, posteriorly a row of shorter setae running parallel to the former; posterior operculum: 11 marginal acuminate setae each side, closely spaced, projecting into lower part of cavity and converging centrally to enclose cavity in kind of grill, posteriorly, a row of widely-spaced setae extending beyond operculum margin; posterior to the former a row of 4 shorter setae behind which is one very long seta (Fig. 2K). Female genitalia: strongly sclerotic with pronounced thicken- ings; thin irregularly-shaped transverse median cribriform plate; numerous minute platelets scattered within sclerotization; spiracles obliquely situated with well defined stigmatic plates (3 setae each) and characteristic for family (Fig. 4W) spermathecae absent. Chaetotaxy: anterior operculum with 6-8 centrally located setae converging to form a pyramid; posterior operculum with 15-17 very short setae on posterior border (Fig. 2L). Sternal chaetotaxy 0° 4-8 weakly divided, 0:18-20: 23-25: 12-14: 18-20: 16-20: 16-18: 14-16: 14-16: 12-14: 10-12: 2, sternites 4-12 uniseriate; 9 4-8 weakly divided, 0: 16- 18: 12-14: 8-10: 14-16: 12-16: 14-16: 14-18: 14-15: 12-14: 10-12: 2, sternites 3-12 uniseriate. Pretarsus: claws obliquely ridged, curving over a trumpet-shaped arolium with delicate serrate rim (Fig. 1H). Heterotarsate. Dimensions (mm): body length 1.-1.16 (1.18-1.51); pedipalps: trochanter 0.14-0.16/0.09-0.10 (0.12-0.18/0.09-0.12), femur 0.37-0.45/0.09- 0.12 (0.41-0.52/0.10-0.12), tibia 0.19-0.27/0,10-0.12 (0.21-0.27/0.10-0.12), chela (with PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 C.M. A. KENNEDY 293 eA Ay UN AA 0-5mm Fig. 3. Pycnodithella harveyi n. sp., male paratype: N, leg I; O, leg IV; P, carapace, dorsal aspect — inset a, epistome not clearly defined in light microscopy) traced from scanning electron micrograph x 1700; Q, left pedipalp, dorsal aspect; R, right chela, lateral aspect; S, right chelicera, dorsal aspect. pedicel) 0.59-0.64/0.12-0.15 (0.64-0.73/0.14-0.18), chela (without pedicel) 0.55-0.60/0.12- 0.15 (0.60-0.70/0.14-0.18), movable finger length 0.30-0.36 (0.36-0.43); chelicera 0.19- 0.25/0.10-0.12 (0.18-0.27/0.12-0.16), movable finger length 0.12-0.14 (0.14-0.18); carapace 0.36-0.43/0.34-0.39 (0.37-0.47/0.39-0.48); anterior eye 0.052-0.054 (0.052-0.056), posterior eye 0.046-0.048 (0.048-0.05); leg I: coxa width 0.09-0.14 (0.09-0.14), trochanter PROG. LINN. SOC. N.S.W., 110 (3), (1988) 1989 294 NEW AUSTRALIAN PSEUDOSCORPION Fig. 4. Pycnodithella harveyi n. sp., paratypes: T, male genitalia, ventral aspect; U, sclerotic, perforated plate designated ain V. V male genitalia and associated sternites; W, female genitalia and associated sternites. 0.09-0.12/0.07-0.10 (0.10-0.18/0.09-0.10), basifemur 0.18-0.23/0.03-0.05 (0.10-0.27/0.03- 0.05), telofemur 0.10-0.18/0.03-0.05 (0.10-0.18/0.03-0.05), tibia 0.09-0.14/0.03-0.04 (0.10-0.16/0.03-0.05), miotarsus 0.16-0.21/0.03 (0.12-0.19/0.02-0.03); leg IV: coxa width PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 C. M. A. KENNEDY 295 0.09-0.10 (0.10-0.12), trochanter 0.10-0.16/0.09-0.11 (0.15-0.91/0.09-0.12), basifemur 0.16- 0.18/0.07-0.12 (0.12-0.27/0.09-0.12), telofemur 0.18-0.21/0.07-0.09 (0.18-0.21/0.07-0.12), tibia 0.16-0.21/0.06-0.08 (0.12-0.27/0.07-0.09), metatarsus 0.09-0.14/0.03-0.05 (0.12- 0.15/0.03-0.05), telotarus 0.18-0.21/0.03-0.04 (0.18'0.21/0.03). Tritonymphs. Pedipapal trochanter 1.0-1.70, femur 3.10-4.57, tibia 1.33-1.80, chela (with pedicel) 3.50-4.60, chela (without pedicel) 3.25-4.10 x longer than broad. Fixed finger with 7 tricobothria, zsb absent, movable finger with 3 trichobothria, 6 absent. Chelicera with 5 simple galea (Fig. 2M), serrula exterior of chelicera with 13-14 lamellae. Cara- pace with 4: 10: 25-35 setae; 0.82-1.0 x longer than broad. ‘Tergal chaetotaxy: 10-12: 10- 12: 10-12: 8-10: 8-10: 8-10: 8-10: 8-10: 8-10: 8-10: 4-6: 2. Sternal chaetotaxy: 8-10: 8-10: 8: 7-8: 8: 8-10: 8-10: 8-10: 7-8: 5-8: 2-3: 2. Coxal chaetotaxy: 1st pedal coxae with mesial process, 1st and 2nd coxae each with 4 medial coxal spines, 0: 4-5: 3, 0: 4-5: 3, 0: 4-5: 7, 0216-82 7-8: Dimensions (mm): body length 0.90-0.99; pedipalps: trochanter 0-10-0.14/0.07-0.09, femur 0.27-0.36/0.07-0.09, tibia 0.12-0.18/0.09-0.10, chela (with pedicel) 0.41-0.52/0.1- 0.14, chela (without pedicel) 0.39-0.50/0.10-0.14, movable chelal finger 0.21-0.32; cara- pace 0.30-0.36/0.32-0.39. Deutonymphs. Pedipalpal trochanter 1.0-1.28, femur 2.57-3.85, tibia 1.28-1.77, chela (with pedicel) 4.0-4.55, chela (without pedicel) 3.90-4.55 x longer than broad. Fixed finger with 6 trichobothria zs¢ and tsb absent, movable finger with 2 trichobothria, sb and b absent. Chelicerae with 4 simple galea. Serrula exterior with 11-12 lamellae. Carapace 3: 8: 20-30 setae, 0.87-0.10 x longer than broad. Tergal chaetotaxy: 8-10: 6-8: 6-8: 8-10: 8- NO GO-O39 Vino-O24-0;4-95 35-472. Stermalichactotaxy-l-2: 1-2:19-6: 9-0: 9-0:.0-079-6; 0-6: 4-5: 4-5: 3-4: 2. Coxal chaetotaxy: Ist pedal coxae with mesial process. Ist and 2nd pedal coxacieachiwith 4 short: medial coxallspines, 0: 2-4: I, 023-42 2,0: 2-4:2510: 2-453. Dimensions (mm): body length 0.70-0.73. Pedipalps: trochanter 0.07-0.09/0.06-0.08, femur 0.18'0.27/0.07-0.08, tibia 0.09-0.16/0.07-0.09, chela (with pedicel) 0.37-0.43/0.09- 0.10, chela (without pedicel) 0.36-0.41/0.09-0.10, movable finger length 0.19-0.25; cara- pace 0.23-0.30/0.25-0.32. Protonymphs. Present indications suggest there is no free living protonymph stage. The question as to whether the embryonic protonymph stage follows a pattern of develop- ment similar to that of Chthonius tetrachelatus Preyssler (Weygoldt, 1968a,b), or if indeed this stage is suppressed in the Australian species, 1s at present under investigation. Comparisons: P. harveyi n. sp. differs from P abyssinica (Beier) 1947 in the sculpture of the carapace which is conspicuously reticulate in contrast to the finely granulated type species. The trichobothria ¢, st, sb in the new species are equidistant (Fig. 1C), as com- pared with the situation in P abyssinica where st of the movable chelal finger is clearly nearer sb than ¢. The new Australian species is also distinctly smaller in most respects than P. abyssinica and has fewer teeth in both chelal fingers. ACKNOWLEDGEMENTS I wish to express my thanks to Dr P. D. Hillyard (British Museum (Natural History), London) for the loan of type material, to Dr M. S. Harlvey (Museum of Victoria, Melbourne) for his guidance and comments offered on the manuscript and to Mr R. J. Oldfield, Macquarie University, for advice in respect to photographic layout. PROC. LINN. SOG. N.S.W., 110 (3), (1988) 1989 296 NEW AUSTRALIAN PSEUDOSCORPION References BALZAN, L., 1891 — Voyage de M. E. Simon au Venezuela: Chernetes (Pseudoscorpiones). Ann. Soc. entomol. France, 60: 504-505, 509. BEIER, M., 1944 — Uber Pseudoscorpioniden aus Ostafrika. Eos Madr., 20: 173-212. ——, 1947 — Zur Kenntnis der Pseudoscorpioniden-fauna des stidlichen Afrika, insbesonders der sud- westund Trockengebiete. Kos Madr, 23: 285-339. CHAMBERLIN, J. C., 1931 — The arachnid order Chelonethida. Stanford Univ. Publs Biol. Sci. 7: 1-284. , and CHAMBERLIN, R. V., 1945 — The genera and species of the Tridenchthoniidae (Dithidae): a family of the arachnid order Chelonethida. Bull. Univ. Utah 35(23), Biol. ser. 9(2). WEYGOLDT, P., 1968a — Vergleichend-embryologische Untersuchungen an Pseudoscorpionen IV. Zeits. fir Morph. der Tiere 63: 111-154. ——,, 1986b — The Biology of Pseudoscorpions. Cambridge, Mass.: Harvard Univ. Press. PROC. LINN. SOC. N.S.W., 110 (3), (1988) 1989 PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES VOLUME 110 NUMBER 4 whe. an te NS eee sins A Study of the Crustacean Zooplankton of Six Floodplain Waterbodies of the Lower Hunter Valley, N.S.W. B. V. TIMMS TIMMS, B. V. A study of the crustacean zooplankton of six floodplain waterbodies of the lower Hunter Valley, N.S.W. Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 297-306. Thirteen species of eulimnetic entomostracans occurred in the waterbodies with Boeckella fluvialis, Calamoecia lucasi, Mesocyclops albicans, Daphnia carinata and Moina micrura dominant. Momentary species composition averaged 1.8 calanoid species, 1.4 cyclopoid copepod and 2.8 cladoceran species. Numerically, calanoids and cladocerans domi- nated (42% each). The low number of cladoceran species and the importance of cala- noids are unusual in a world context, so possible reasons for these are explored. Most differences between lagoons are explained in terms of relative habitat stability and the presence/absence of predators. Seasonal variation in total numbers was erratic. All copepods were perennial, Daphnia carinata, D. lumholtzi and Bosmina meridionalis were almost perennial, while the remaining cladocerans were distinctly seasonal. Members of the congeneric pairs Diaphanosoma excisum— D. unguiculatum, Daphnia carinata— D, lumholtzi, and Ceriodaphnia cornuta— C. dubia’ peaked at different times. Four of the waterbodies dried periodically, but zooplankton populations recovered quickly on refilling, much faster than the colonization of new sites. Successional sequences depended on season, but generally cyclopoids and Moina micrura were fast colonizers, followed by Boeckella fluvialis, Daphnia carinata, then Bosmina meridionalis, and lastly Calamoecia lucasi and Daphnia lumholtzi. B. V. Timms, Sciences Department, Avondale C.A.E., PO. Box 19, Cooranbong, Australia 2265, manuscript received 4 August 1987, accepted for publication 17 February 1988. KEY WORDS: Calanoida, Cyclopoida, Cladocera, seasonal distribution, community structure, plankton predators, colonization. INTRODUCTION Compared with other aquatic communities, the crustacean zooplankton of Australian lakes is relatively well known. There is information on taxonomy and distri- bution of the major components (e.g. Bayly, 1961, 1964, 1966, 1979, on calanoid cope- pods; Bayly and Morton, 1978, Morton, 1985, on some cyclopoid copepods; Smirnov and Timms, 1983, on cladocerans), and on the association of species in various areas (Mitchell, 1986; Shiel et a/., 1982; Timms and Morton, 1988). Study continues on the value and methodology of conceptualizing associations and on the relative importance of physicochemical and biological factors in determining species composition and seasonal change. Earlier work used and misused the statistic ‘momentary species com- position and concentrated on the influence of physiocochemical parameters (e.g. Timms, 1970a), though the role of competition was not completely ignored (e.g. Bayly, 1966: Jolly, 1966). More recently the effect of vertebrate and invertebrate predation has been highlighted (Geddes, 1986; Grant and Bayly, 1981; Reynolds and Geddes, 1984) and new methods devised to depict species composition (Mitchell, 1986). Most of these interpretations have been based on spot sampling or on short-term studies with some narrow goal in mind. An alternative approach is to monitor plankton over many years and to interpret general trends. Such studies, e.g. Jolly (1966) on the Sydney reservoirs, lack the rigour of specific goal short-term studies, but they can pro- vide useful data. For instance, long-term studies are more likely to record all species at a given site and provide a more accurate assessment of the ‘average’ plankton community. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 298 CRUSTACEAN ZOOPLANKTON Also such studies are able to record in proper context the effect of irregular events such as floods and droughts. In this paper the crustacean zooplankton of six small floodplain waterbodies of the lower Hunter Valley have been studied monthly for just over 5 years. This period en- compassed two droughts and a number of minor floods. With the broadscale distri- bution of crustacean zooplankters in the region known (Timms, 1970a) and comparative seasonal data available for nearby reservoirs (Timms, 1970b), the aim of this study is to document the long-term composition and variability of plankton in natural sites in mid-eastern Australia and to note the major influencing factors. STUDY SITES AND METHODS Six floodplain waterbodies of the lower Hunter River near Maitland were sampled monthly from October, 1979 to December 1984. Geomorphically they are blocked valley lakes, but locally they are called ‘lagoons’, a descriptor used in Australia for any small in- land waterbody, so this practice is adopted here. Their location, geomorphic parameters and salient physicochemical features are given in Timms (1987b). Lagoons la and 1b contained water throughout the study, and Lagoon 3 was nearly permanent, only drying for 3 out of the 63 months. The other lagoons were intermittently present — Lagoon 2 was dry for a total of 27 months, Lagoon 4 for 26 months and Lagoon 5 for 31 months (see Timms, 1987b: fig. 4). All lagoons filled from their local catchment. On occasions macrophytes grew too extensively to allow open water sampling in Lagoon 3 and less frequently in Lagoon 2. On each sampling occasion, a conical net 30.5cm in diameter and mesh size 159um was used to collect zooplankton from open water. If the site was > 40cm deep an oblique haul from near the bottom to the surface was taken from a boat, but if the depth was < 40cm deep a horizontal tow was taken by wading through the lagoon. In both cases the net was towed at approximately 20cm sec’ for 2 minutes over 25m, thus filtering 1820L at most, but almost certainly much less (Bottrell et al., 1976). The method is semi- quantitative at best, but consistent. Samples were preserved in 5% formalin. Sub- samples were taken from each collection so that about 1000 individuals were enumer- ated. Plankters were assigned to species, though this was incomplete for cyclopoid copepods (small Mesocyclops albicans were included with Thermocyclops decipens and the latter were lumped with Eucyclops spp.). Where two species of calanoid copepod co- occurred, copepodites were included in species totals according to the ratio of adult individuals identified for the species present. RESULTS (a) Species Composition and Community Structure Altogether 13 eulimnetic entomostracans, comprising 3 calanoid copepods, 2 cyclopoid copepods and 8 cladocerans occurred in the 6 lagoons (Table 1). Eight littoral species (2 cyclopoids, 2 ostracods, 3 chydorid cladocerans and Simocephalus) sometimes were collected from open waters, but generally in small numbers. Eucyclops spp. may have occurred in lagoons other than No. 3, but it is consistent they should be found in the site with the best developed littoral macrophytes. In addition shrimp larvae (? Paratya australiensis) occurred in Lagoons la, 1b, and 3, mainly in summer, and a number of rotifers, including Brachionus spp., Keratella sp. and Asplanchna sp., were common in summer-autumn. Species composition was similar, but not identical, in each lagoon (Table 1). The presence of possible plankton predators was also noted. European Carp (Cypr- nus carpio 1.) was common in Lagoon 1b and colonized Lagoon 1a during 1983, but were PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 B. V. TIMMS 299 TABLE 1 Relative Importance of Entomostracan Zooplankters in the Lagoons Lagoons Species la Ib 2 3 4 5 1. Duaphanosoma excisum Sars Dee 4.3 1.3 Apo 1.6 = 2. Diaphanosoma unguiculatum Gurney 2.8 Dy 0.4 0.6 2.6 189 3. Moina micrura Kurz 9.8 24: eZ 4.7 9.5 eZ 4. Bosmina meridionalis Sars Za) Usd SEA. 22 4.4 0.7 5. Daphnia carinata King 14.0 3.6 23.4 6.6 2233 28.0 6. Daphnia lumholtzi Sars 2.8 DY = 223 — — 7. Ceriodaphnia cornuta Sars 3.6 3.6 0.5 33 0.3 0.6 8. Certodaphnia dubia’ Richard De? De2 2 5e3 6.7 1.0 9. Chydorids+ 0.2 0.4 1.7 11 0.3 0.7 10. Sumocephalus vetulus elisabethae (King) = = 0.2 = = - 11. Newnhamia fenestrata King = = 2.0 — — = 12. Cyprinotus fuscus Henry = = 0.8 = 0.2 1.6 13. Boeckella fluvialis Henry 12.0 8.4 18.7 21.5 14.4 el) 14. Boeckella triarticulata Thomson = = = — — 1.4 15. Calamoecia lucas: Brady 28.1 30.0 17.6 38.7 23) 10.4 16. Mesocyclops albicans (Smith) 12.0 10.3 15.0 6.4 12.4 11.8 17. Thermocyclops decipiens (Kiefer) 6.2 6.1 Hel Dap 5.0 18. Eucyclops sp. nov. = = = , 3.0 = = 19. Eucyclops sp. prob. ewacanthus (Sars) — = = = = * Calculated by averaging percent composition for each month in each lagoon. + includes Chydorus sphaericus s.J. O. F. Muller, Dunhevedia crassa King, and Alona diaphana King. still not common by December 1984. Mosquito Fish (Gambusza affinis Baird and Girard) were numerous in Lagoons la, 1b and 3. Notonectids (Anzsops spp.,) occurred in all lagoons, particularly in Lagoons 2, 4 and 5, and were most common November to April (Fig. 2). No Chaoborus were ever caught in plankton tows or in a few Ekman grab samples taken in a benthic study (author, unpublished). The relative importance of species varied between sites (Table 1). The main differ- ences were between the permanent to near-permanent lagoons (Nos. la, 1b, 3) which also had fish and the intermittently-present lagoons (Nos. 2,4, 5) which lacked fish. Daphnia lumholtzi was absent from the latter group and Calamoecia lucas, Diaphanosoma excisum, Certodaphnia cornuta, were less common in them, though the differences may not be significant given the small number of sites. Three species were restricted to these intermittently present lagoons — the ostracods Cyprinotus fuscus found in all three and Newnhamia fenestrata in Lagoon 2 only, and Boeckella triarticulata in Lagoon 5 for just 3 months. On the other hand Daphnia carinata was most common in these lagoons. It and Boeckella fluvialis were least important in Lagoon 1b in which carp were abundant. On average the six lagoons contain 1.8 calanoid species, 1.4 cyclopoid species and 2.8 cladoceran species (Table 2). There is seasonal variation in these averages, but it is erratic both between sites (Table 2) and between years (Fig. 1), especially for clado- cerans. Of the three groups, however, cladocerans are more consistently present in per- manent rather than intermittently present lagoons (t = 9.5, P < 0.001, DF = 4). (b) Seasonality Total numbers of zooplankton per month in each lagoon fluctuated ca10-100 fold with little apparent seasonal regularity (Fig. 1), but when data were averaged for each month in each lagoon, some general trends emerged (Fig. 2). In all lagoons numbers tended to be lowest in winter and highest in autumn. The two permanent lagoons (la, PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 too weedy a 7 < too weedy too weedy too weedy é 2 Fig 1. Relative abundance and approximate total numbers per standardized collection of crustacean zooplankton in the six lagoons (1a, 1b, 2, 3, 4, 5) for the period October 1979 to December 1984. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 B. V. TIMMS 301 D excisum it D carinata B fluvialis A M albicans D unguiculatum _ aad lumholtzi B MERE enietaln ah. cornuta other cyclopoids M micrura C fuscus Ceriodaphnia spp C lucasi 6 backswimmers B meridionalis chydorids 400 40 T O 20 ES An iAy On D CMIV Fig. 2. Seasonal variation in the cumulative monthly importance values (CMIV) of various crustacean species and notonectids for the period October, 1979 to December, 1984. These are derived by giving each monthly occurrence of each species in each lagoon a numerical value (1 = when relative abundance < 2.5%, 2 = 2.5- 10%; 3 = 10-25%; 4 = 25-50%; 5 = >50%) and then summing. Also shown is the average number of zooplankton per standardized collection for each month in each lagoon. 1b) had elevated numbers during October-November and in April. The intermittently- present lagoons (2, 4, 5) had only one major peak in April-June, and the near- permanent lagoon (3) had an intermediate pattern. Kite diagrams of the percentage abundance of each species over the 63 month study period (Fig. 1) and histograms of cumulative monthly importance values (Fig. 2) show various patterns of seasonality. The calanoids Boeckella fluvialis and Calamoecia lucast were perennial, with a period of minimal importance in summer-early autumn, Le. December to March in B, fluvialis and January-April in C. lucasi. Boeckella triarticulata oc- curred only from August to October which does not indicate its full seasonal capabili- ties, as this occurrence is believed to be a failed colonization attempt (see later). Both Mesocyclops albicans and Thermocyclops/Eucyclops spp. were perennial or almost so (Fig. 2), but with distinct peaks from February to May and from September to February respec- tively. Three of the 8 eulimnetic cladocerans, Daphnia carinata, D. lumholtzi and Bosmina meridtonalis may have been perennial (Fig. 2), though they were not continuously present in any lagoon (Fig. 1). Daphnia carinata peaked seasonally in May-October and B. meri- dionalis in April-July. The other 5 species were distinctly seasonal (Fig. 2) — Diaphano- soma excisum from January to May, D. unguiculatum from July to November, Moina micrura from November to May, Certodaphnia cornuta from December to July, and C. dubia’ from PROC. LINN. SOC. N.S.W,, 110 (4), (1988) 1989 302 CRUSTACEAN ZOOPLANKTON July to October. Fig. 2 suggests some overlap in the presence of each species in the three congeneric pairs of cladocerans, but this rarely happened in individual lagoons (Fig. 1). Littoral cladocerans tended to stray into open water in any month, but with a peak in spring, and the ostracod Cyprinotus fuscus was also recorded in almost any month, except in January, February and July. (c) Influence of Floods and Droughts No major river floods occurred during 1979-84, but there were a number of periods of heavy rainfall, local flooding and rapid rises in the levels of the lagoons (Timms, 1987b). No particular effect of these inflows was detected, though in general population peaks occurred in April-May, two months after peak seasonal rainfall. The lagoons refilled after being dry on 7 occasions, twice each in Lagoons 2, 4 and 5 and once in Lagoon 3 (Fig. 1). Generally the order of response was Moina micrura > Mesocyclops albicans > Boeckella fluvialis < Daphnia carinata. On one occasion each Bosmina meridionalis and Diaphanosoma excisum reestablished quickly. Boeckella fluvialis typically established sizeable populations a month or so earlier than Calamoecia lucast. DISCUSSION (a) Species Composition Almost all of the species recorded in the six study sites are within their known distribution in northeastern N.S.W. (Smirnov and Timms, 1983; Timms, 1970a). The only exception is Ceriodaphnia dubia’ but re-examination of most of the collections used for Timms (1970a) and (1970b) showed it indeed occurred in a few. More notable are some omissions. Boeckella minuta is absent, though common in reservoirs and some natural lakes in mid-eastern Australia, thus confirming its preference for newly-created habitat (Bayly, 1979; Timms, 1970a). Gladioferens spinosus is also absent, despite being present in the nearby Hunter River and in local reservoirs which draw their water from it (Timms, 1970b). It seems this species is dispersed almost exclusively through the aquatic medium (Timms, 1970b, 1973), but apparently conditions during major floods when the lagoons are in contact with the Hunter River are unsuitable for dispersal for other reasons. Of the factors likely to affect species composition and seasonal distribution of crustacean zooplankton, some data are available on the degree of permanence of the lagoons (the major physicochemical difference between them — Timms, 1987b) and on the presence of fish and notonectids. Many differences between the 3 permanent and 3 intermittently-present lagoons (Table 1) can be explained by (a) the relatively poor colonization and recovery ability of Calamoecia lucast (Maly, 1984a; Timms, 1970a, 1987a) and also of Daphnia lumholtzi (Timms, 1970a, 1987a) and hence their lesser importance (C. lucasz) or absence (D. lumholtzi) from the three intermittently present lagoons, and (b) the lagoons being dry when species such as Diaphanosoma excisum, D. unguiculatum and Ceriodaphnia cornuta are normally abundant in lagoons and reservoirs in the area (Fig. 1 and Timms, 1970b). While no specific data are available on predation by fish in these lagoons, the relatively poor performance of the two largest species, Daphnia carinata and Boeckella fluvialis in the two lagoons with fish is consistent with the work by Geddes (1986) on zooplankton composition in farm dams, Hume et al. (1983) and Straskraba (1965) on carp diets and Lloyd (1986) on the influence of mosquito fish on zooplankton. Also the abundance of predatory notonectids in the lagoons from November to April probably has profound effects on abundances of many species (cf, Geddes, 1986). The decline in early summer of Daphnia carinata and possibly also of other large species such as D. PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 B. V. TIMMS 303 lumholtzt, Diaphanosoma unguiculatum and Boeckella fluvialis (Figs 1 and 2) could be explained by predation pressure by notonectids. Seasonal changes in the relative abun- dance of the smaller species, Diaphanosoma excisum, Moina micrura, Ceriodaphnia spp. and Calamoecia lucas: do not coincide with the abundance of notonectids (Fig. 2). (b) Seasonal variation Erratic seasonal variation in zooplankton numbers as seen in these lagoons is to be expected in small lakes (Pennak, 1949), though there is a trend towards late spring and autumn maxima as seen in larger reservoirs nearby (Timms, 1970b), The perennial occurrence of Boeckella fluvialis, Calamoecia lucast, Mesocyclops albicans (formerly misidenti- fied as M. leuckarti in Australia), Daphnia carinata and Bosmina meridionalis is consistent with results of other studies in southeastern Australia (Geddes, 1968, 1984: Jolly, 1966; Timms, 1970b: Walker and Hillman, 1977). Daphnia lumholtzi is almost perennial in these lagoons, but with a distinct minimal in summer, whereas in nearby reservoirs (Timms, 1970b) it was most abundant in summer and usually absent in winter. Its absence from the 3 shallower ephemeral lagoons studied here and from nearby reser- voirs during low water levels (Timms, 1970b), suggests that other factors associated with low water levels may inhibit its development. The seasonal distribution of other species mirror those reported elsewhere in southeastern Australia. Diaphanosoma excisum is abun- dant in summer-early autumn (Timms, 1967, 1970b), D. unguzculatum is a spring species (Geddes, 1984; Shiel et al., 1982; Walker and Hillman, 1977), Mona micrura blooms in summer (Geddes, 1968, 1984; Shiel et a/., 1982; Timms, 1970b) and Ceriodaphnia cornuta is also a summer species (Timms, 1970b). No comparative data are available for Cerzo- daphnia dubia: All species, perennial and seasonal, appear to be multivoltine. (c) Community Structure Zooplankton community structure is often described by the parameter momentary species composition (MSC), which is the number of species present at any one time (Pennak, 1957). Mitchell (1986) has challenged its value and validity, but most of the in- adequacies are due to limited data bases. In the present lagoons this is not a problem as MSCs are based on many observations totalling 32-63 according to lagoon. The average MSC in the six lagoons (Table 2) is within the range reported for lakes and ponds in southern Australia, though on a world comparative basis the number cladocerans is lower in these lagoons and in Australian sites in general (Mitchell, 1986, Timms, 1970a). In that cladoceran assemblages are less rich in the study lagoons which are intermittently present, perhaps the ephemeral status of many Australian sites con- tributes to this Australia versus World difference. The explanation for this lies in the dis- tinct seasonality of most cladocerans and hence greater likelihood of their absence from sites which dry intermittently. This also promotes erratic variation in species richness among cladocerans whereas among calanoids community structure is more stable (Table 2). Further study on this factor as well as others mentioned by Mitchell (1986) is needed. A further method of comparing assemblages is by calculating the average percentage composition of the component species from regular samples taken over at least a year. This is done in Table 1 and has already been used (see above) to highlight similarities and differences between lagoons. Also of interest is the relative overall contribution by cladocerans (42%), calanoids (42%) and cyclopoids (16%) (Table 1). These proportions are not dissimilar from those (36 : 49 : 15 respectively) for Australian lakes reviewed by Mitchell (1986) and provide further evidence for the relatively greater contribution by calanoids in Australia vis-a-vis most other countries. Possible reasons for this are partly explored by Mitchell (1986), to which perhaps should be added the apparently great PROG. LINN. SOC. N.S.W., 110 (4), (1988) 1989 304 CRUSTACEAN ZOOPLANKTON TABLE 2 Average Momentary Species Composition in the Lagoons Calanoida adaptability and comparatively good dispersal powers of many centropagid copepods in Australia (Bayly, 1964, Maly, 1984a, Timms, 1970a, 1987a). (d) Recovery from Dryness The succession of species following refilling is broadly similar to that seen in the colonization of an entirely new waterbody (Timms, 1987a). Moina micrura and cyclo- poids are fast colonizers, followed by Boeckella fluvialis and Daphnia carinata, then Bosmina meridionalis and finally, Calamoecia lucasi and Daphnia lumholtz1. The exact sequence in any succession is influenced by season. In the present lagoons, if the February and April recoveries are compared, Daphnia carinata did better and Moina micrura worse in April in accordance with their usual importance in these months (Figs 1 and 2). Also for similar reasons, Diaphanosoma excisum did well in one February recovery and Bosmina meridionalis in one April recovery. Generally cyclopoids and cladocerans responded quicker to refill- ing than calanoids, which is explained by the shorter life cycles of the former (Hutchin- son, 1967). Of the calanoids, Boeckella fluvialis typically responded quicker than Calamoecia lucas: to refilling, as it produces more eggs and hence more offspring (Maly, 1984a). There are two significant differences between the colonization of a new waterbody and recovery in a refilled one. The first is the absence of Boeckella minuta, a prime colonizer of new habitats (Bayly, 1979; Maly, 1984a; Timms, 1970a, 1987a). Perhaps it did arrive during recovery (and at other times as well) but was reproductively swamped or competitively excluded by the incumbent Boeckella fluvialis (Maly, 1984b). Similar reasons probably account for the failure of B. triarticulata to establish in Lagoon 5. The PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 B. V. TIMMS 305 other difference is the greater rapidity of succession in lagoons recovering from dryness. This could be caused by the greater numbers of resting eggs (and hence nauplii hatch- ing) in the recovery lagoon compared with the presumed few dissemules arriving at a new waterbody. ACKNOWLEDGEMENTS I wish to thank Dr Patrick De Deckker for identifying ostracods, Dave Morton for identifying cyclopoids , and Dr Ian Bayly and Dr Rus Shiels for their comments on the manuscript. References BayLy, I. A. E., 1961. — A revision of the inland water genus Calamoecia (Copepoda: Calanoida). Aust. J. Mar. Freshw. Res. 12: 54-91. — 1964. — A revision of the Australian species of the freshwater genera Boeckella and Hemiboeckella (Copepoda: Calanoida). Aust. J. Mar. Freshw. Res. 15: 180-238. — 1966. — The Australian species of Diaptomus (Copepoda: Calanoida) and their distribution. Aust. /. Mar. Fresh. Res. 17: 123-134. — 1979. — Further contributions to knowledge of the Centropagid genera Boeckella, Hemiboeckella and Calamoecia (Athalassic calanoid copepods). Aust. J. Mar. Freshw. Res. 30: 103-127. and Morton, D. W., 1978. — Aspects of the zoogeography of Australian microcrustaceans. Verh. Int. Ver. Limnol. 20: 2537-2540. BOTTRELL, J., et al., 1976. — A review of some problems in zooplankton production studies. Norw. J. Zool. 24: 419-456. GEDDES, M. C., 1968. — Studies on the Marshall Reserve pond with special reference to Boeckella triarticulata Thompson (Copepoda: Calanoida). Clayton, Victoria: Monash University, honours thesis unpubl. —— 1984. — Seasonal studies on the zooplankton community of Lake Alexandrina, River Murray, South Australia, and the role of turbidity in determining community structure. Aust. J. Mar. Freshw. Res. 35: 417-426. —— 1986. — Understanding zooplankton communities in farm dams: the importance of predation. Jn DE DECKKER, P., and WILLIAMS, W. D., (eds), Limnology in Australia: 387-401. Melbourne/Dordrecht: CSIRO/Junk. GRANT, J. W. G., and Bay Ly, I. A. E., 1981. — Predator induction of crests in morphs of the Daphnia carinata King complex. Limnol. Oceanogr. 26: 201-218. Hume, D. J., FLETCHER, A.’R., and Morison, A. K. 1983. — Final Report. Carp Program. Arthur Rylah Inst. for Environ. Res., Fish. and Wildl. Div., Min. Cons., Victoria. 213 pp. HUTCHINSON, G. E., 1967. — Treatise on Limnology. Vol. II. Introduction to Lake Biology and the Limnoplankton. New York: John Wiley and Sons. JOLLy, V. H., 1966. — The limnetic crustacea of six reservoirs in the Sydney area of New South Wales. Verh. Int. Ver. Limnol. 16: 727-734. LLoyD, L. N., ARTHINGTON, A. H., and MILTON, D. A. 1986. — The mosquitofish — a valuable mosquito- control agent or a pest? Jn KITCHING, R. J., (ed.), The Ecology of Exotic Animals and Plants in Australia: 7- 25. Brisbane: John Wiley and Sons. MALY, E. J., 1984a. — Dispersal ability and relative abundance of Boeckella and Calamoecia (Copepoda: Calanoida) in Australian and New Zealand waters. Oecologia (Berl. ) 62: 173-181. —— 1984b. — Interspecific copulation in and co-occurrence of similar-sized freshwater centropagid cope- pods. Aust. J. Mar. Freshw. Res. 35: 153-165. MITCHELL, B. D., 1986. — Entomostracan zooplankton communities of Australian freshwater lakes and ponds. Jn DE DECKKER, P., and WILLIAMS, W. D., (eds), Limnology in Australia: 369-386. Melbourne/Dordrecht: CSIRO/Junk. Morton, D. W., 1985. — Revision of the Australian Cyclopoidae (Copepoda: Cyclopoida) I. Acanthocyclops Kiefer, Diacyclops Kiefer and Australocyclops gen. nov. Aust. J. Mar. Freshwat. Res. 36: 615-634. PENNAK, R. W., 1949. — Annual limnological cycles in some Colorado reservoir lakes. Ecol. Monogr. 19: 233-267. —— 1957. — Species composition of limnetic zooplankton communities. Limnol. Oceanogr. 2: 222-232. REYNOLDS, J. G., and GEDDES, M. C., 1984. — Functional response analysis of size-selective predation by the notonectid predator Anzsops deanei (Brooks) on Daphnia thomson: (Sars). Aust. J. Mar. Freshw. Res. 35: 725-733. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 306 CRUSTACEAN ZOOPLANKTON SHIEL, R. J., WALKER, K. F., and WILLIAMS, W. D., 1982. — Plankton of the lower Murray River, South Australia. Aust. J. Mar. Fresh. Res. 33: 301-327. SMIRNOV, N. N., and Timms, B. V., 1983. — A revision of the Australian Cladocera (Crustacea). Rec. Aust. Mus., Suppl. 1: 1-132. STRASKRABA, M., 1965. — The effect of fish on the number of invertebrates in ponds and streams. Mitt. Int. Verein. Limnol. 13: 106-127. TIMMS, B. V., 1967. — Ecological studies on the Entomostraca of a Queensland pond with special reference to Boeckella minuta Sars (Copepoda: Calanoida). Proc. Roy. Soc. Qd79: 41-70. — 1970a. — Chemical and zooplankton studies on lentic habitats of north-eastern New South Wales. Aust. J. Mar. Freshw. Res. 21: 11-33. — 1970b. — Aspects of the limnology of five small reservoirs in New South Wales. Proc. Linn. Soc. N.S.W. 95: 46-59. —— 1973. — A limnological survey of the freshwater coastal lakes of east Gippsland, Victoria. Aust. J. Mar. Freshw. Res. 24: 1-20. — 1987a. — Colonization of two large farm dams on the central coast of New South Wales by crustacean zooplankton. Bull. Aust. Soc. Limnol. 11: 9-14. — 1987b. — Geomorphic and physicochemical features of floodplain waterbodies of the lower Hunter Valley, N.S.W. Proc. Linn. Soc. N.S.W. 109: 311-324. and Morton, D. W., 1988. — Crustacean zooplankton assemblages in freshwaters of tropical Aus- tralia. Hydrobiologia 164: 161-169. WALKER, K. F., and HILLMaN, T. J., 1977. — Limnological survey of the River Murray in relation to Albury-Wodonga, 1973-1976. Report on behalf of Gutteridge, Haskins & Davey to the Albury- Wodonga Development Corporation, Albury. PROC. LINN. SOC. N.S.W,, 110 (4), (1988) 1989 Seed Type and Seed Surface Patterns in Calandrinia sens. lat. (Portulacaceae) SYEDA SALEHA TAHIR and ROGER CAROLIN SYEDA SALEHA TAHIR, & CAROLIN, ROGER, Seed type and seed surface patterns in Calandrinia sens. lat. (Portulacaceae). Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 307-316. Seed types of Calandrinia are defined as six shapes which are determined by the shape of the embryo and the amount of perisperm present. Ten surface patterns are described and the presence or absence of an aril are considered. The species usually have characteristic shapes and surface patterns but these features rarely characterize the segregate genera or the sections. Within some of the segregate genera and sections, however, there appear to be definite, and sometimes characteristic, phylogenetic trends in these features. Syeda Saleha Tahir and Roger Carolin, John Ray Herbarium, University of Sydney, Australia 2006; (Syeda Saleha Tahir, new address, Botany Department, University of Sind, Jamshoro, Sind, Pakistan); manuscript received 14 September 1987, accepted for publication 23 March 1988. INTRODUCTION Seed type and seed surface pattern have been used extensively in plant taxonomy. Carolin (1987) has shown that the former, in particular, is useful at the generic level in the family Portulacaceae and has indicated that in certain genera, surface pattern can be useful in distinguishing between species (see also Cullen, 1953). Furthermore, the aril or strophiole, in this case derived from the funicle, has been used in the past to discriminate between genera, e.g., Pax and Hoffman (1935). This investigation seeks to establish the use of these features for the taxonomy within the genus Calandrinia. The genus Calandrinia consists of about 130 species of which about 50 occur in Australia. Carolin (1987) has suggested that the genus be divided into five genera, Calan- drinia sens. strict., Cistanthe, Baitaria, Schreiterra and Rumicastrum. The necessary combi- nations have not been made, except in Schreiteria, and here they will all be referred to as Calandrinia with the segregate name in parentheses. Although seed surface pattern has been used in the past to distinguish between some Australian species, e.g., Black (1948) and Reiche (1888), the only general survey of either seed type or surface pattern in these species is that of Syeda (1980). This contribution details the variation in these charac- ters, provides a terminology and suggests a phylogeny for these characters. MATERIAL AND METHODS The mature seeds were taken from herbarium specimens and mounted on a stub with double sided sticky tape. They were then coated with 200-400A thickness of gold in a polaron coating machine and examined with a JSM-U3 scanning electron micro- scope. Iwo to three samples from different collections were examined for 77 species in this manner but in most cases only one voucher is listed here (Table 1). Where available, the seeds of most specimens of each species were examined using a stereo-microscope to check for variation within a species. Hand sections were cut to ascertain the shape of the embryo and the distribution of the perisperm. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 308 SEED TYPES OF CALANDRINIA A. C. strophiolata C. C. tenuifolia D. C. grandiflora E. C. compressa Fig. 1. A. C. strophiolata; B. C. polyandra; C. C. tenuifolia; D. C. grandiflora; E. C. compressa. (Bars indicate 100). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 SYEDA SALEHA TAHIR AND ROGER CAROLIN 309 RESULTS SEED SHAPES The seed shape presented some problem since the differences between species, though often clear, are relatively slight and difficult to describe in words. The shapes defined in the chart of plane shapes reproduced in Stearn (1966) are not entirely satis- factory for 3-dimensional objects. Nevertheless, many workers do use them in this con- text, usually apparently referring to the projected shadow across the long axis. The shapes of Calandrinia seeds, however, do not always correspond to the standards even when thus projected onto a plane. Stearn also provides a chart of spore shapes from Ainsworth and Bisby; where these terms apply we have used them. Moreover some of the shapes integrade with each other and the decision as to their classification in some cases is a little arbitrary. The terms are redefined below in this particular context and illustrated in Figs 1 and 2. Seed shape is a function of the shape of the embryo and, to a lesser extent, the amount of perisperm. The curvature and length of the embryo, and the relative growth of the radicle in relation to the cotyledons, determine the position of the funicle. The schematic diagrams in Fig. 3 illustrate this: Oblong/Orbicular: Embryo almost completely encircling the perisperm; funicle attach- ment symmetrical at the base; + compressed. The two shapes intergrade closely, e.g. C. oblonga (Fig. 2F). See Fig. 3A, B. Notched Oblong/Orbicular: As for oblong but growth of both radicle and cotyledons continued below the funicular attachment, giving a basal notch, e.g. C. tenuzfolia (Fig. 1C), C. polyandra (Fig. 1B). See Fig. 3C. Obovoid to globular: Embryo almost encircling the perisperm but growth of the radicle continued below the funicular attachment thus causing the latter to be asymmetric; sometimes this distortion becomes extreme; not or very little compressed, e.g. C. ptycho- sperma (Fig. 2C), C. reticulata (Fig. 2D), C. grandiflora (Fig. 1D). See Fig. 3D. Asymmetric clavate: Embryo not quite encircling the perisperm, funicle attached to one side of a basal point; not compressed, e.g. C. lehmanni., This represents a type inter- mediate between obovoid/globular and curved. Curved: Embryo not encircling perisperm but curved distally; funicle attached to one side of a basal point; not compressed, e.g. C. primuliflora (Fig. 2E). See Fig. 3F. Narrow-obovoid: A very narrow seed but similar to the obovoid form described above; scarcely compressed, e.g. C. disperma. There are, of course, intermediate states, some of which may indicate phylogenetic series. SEED SURFACE PATTERN Here also the terms described by Stearn (1966) for surface patterns need some amendment as defined below: Smooth: No prominent pattern, the outline of the epidermal cells usually visible but the surface between the radial walls flat, e.g. C. strophiolata (Fig. 1A). Colliculate: The surface of each epidermal cell is slightly raised into a shallow dome forming a closely packed pattern, e.g. C. polyandra (Fig. 1B). Reticulate: The radial walls of each epidermal cell are raised into a ridge, e.g. C. reticulata (Fig. 2D). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 310 SEED TYPES OF CALANDRINIA A. C. arenaria SS E. C. primuliflora Ta F. C. oblonga i Fig. 2. A. C. arenaria; B. C. pickeringii; C. C. ptychosperma; D. C. reticulata; E. C. primuliflora; F. C. oblonga. (Bars indicate 100y). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 SYEDA SALEHA TAHIR AND ROGER CAROLIN lil Fig. 3. Schematic diagrams to show seed shape and its associated features. A. Oblong/(orbicular); B. (Oblong)/orbicular: C. Notched oblong/orbicular; D. Obovoid; E. Narrow obovoid; F. Curved. Papillate: The centre of the outer tangential wall of the epidermal cell is projected into a short rounded papilla e.g. C. compressa (Fig. 1E). Generally the cells of the epidermis tend to be aligned into rows and thus, so do the papillae. In some species this is more pronounced than in others and in this case the pattern may be referred to as aligned- papillate, e.g. C. arenicola. Aculeate: Similar to tuberculate in which the papillae are extended into a long narrow hair-like process, e.g., C. grandiflora (Fig. 1D). Polypiform: In this case the papillae have short lobes over their surface, e.g., C. arenarza (Fig. 2A). Papillate-punctate: Similar to papillate but with a small pit at the corner junctions of each cell, e.g. C. pickeringi (Fig. 2B). Ribbed: The surface is marked by prominent ribs which are the confluent papillae of adjacent aligned cells, e.g. C. ptychosperma (Fig. 2C). Tuberculate: The papillae project into broad finger-like processes e.g. C. lehmannit. Verrucate: Similar to tuberculate but the tubercles are somewhat shorter and, although the top 1s smooth, the sides are ridged as though they were thinner, thus giving a warty appearance, e.g. C. primuliflora (Fig. 2E). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 ob2 SEED TYPES OF CALANDRINIA ARIL It is often difficult to determine whether an aril is present or not since small frag- ments of the funicle may be left on the seed due to the abscission occurring well below the insertion of the funicle on the seed. The fragment left may or may not function as an elaiosome in ant dispersal. It seems possible that a well developed aril is an elaiosome as in C. grandiflora (Carolin, pers. obs.). This true aril is the swollen upper part of the funicle and the primary abscission occurs below it; this is registered as ‘+’ e.g. C., ptycho- sperma (Fig. 2C), C. strophiolata (Fig. 1A), C. grandiflora (Fig. 1D), C. arenaria (Fig. 2A). When only a slight swelling, or none at all, occurs above the primary abscission but a fragment of funicle remains on the seed this is registered as ‘+’ e.g. C. polyandra (Fig. 1B). When the primary abscission occurs at the insertion of the funicle on the seed this is registered as — e.g. C. tenuifolia (Fig. 1C), C. compressa (Fig. 1E), C. primulzflora (Fig. 2E). The results are summarized in Table 1, in which the Australian species are grouped according to the sections of Syeda (1980) and the others are grouped according to Carolin (1988). TABLE 1 Species examined for seed characteristics in the course of this investigation (Vouchers are given using the herbarium number where one is available. Where one is not available the herbarium is indicated and the collector and collector’s number are given. Herbarium codes are taken from Index Herbariorum) Species name shape pattern aril Voucher Calandrinia (Cistanthe) sect. Cistanthe C. arenaria Cham. obov Sm/Co + CONC 47320 C. cachinalensis Phil. obov Po + = NOVOUCHER C. coquimbensis Barn. obov Ac + NO VOUCHER C. frigida Barn. obov Pp + CONC 32814 C. glaucopurpurea Reiche obov Po + | NOVOUCHER C. grandiflora Lindl. obov Ac + Carolin 14022 syp C. litoralis Phil. obov Po + CONC 47322 C. longiscapa Barn. obov Po + NO VOUCHER C. maritima Nutt. obov Ac + NOVOUCHER C. oblongifolia Barn. obov Po + Carolin 14112 syp C. solisi Phil. obov Po + | NOVOUCHER C. thyrsoidea Reiche obov Ac + CONC 8396 Calandrinta (Cistanthe) sect. Amarantoideae C. ambigua Nutt. obov Pp — _B 1417582 C. calycina Phil. ob/or Sm/Co — CONC 33144 C. spicigera Phil. ob/or Sm/Co — CONC 47320 Silvaea pachyphylla Phil. ob/or Sm/Co — CONC 9365 Calandrinia ( Baitaria) sect. Dianthoideae C. andicola Gill. ex Arn. n.ob/or Sm — CONC 49244 C. cistiflora Gill.ex.Arn. n.ob/or Co — CONC 3200 C. gayana Barn. n.ob/or Co/Sm — CONC 4898 C. patagonica Speg. ob/or Co — CONC 34827 C. tenuifolia Phil. n.ob/or Co — CONC 32104 Calandrinia sens. strict. C. axilliflora Barn. obov Pp + CONC 49732 C. compressa Schrad. obov Pp/Co + Carolin 14000 syp C. cthata (Ruiz & Pavon) DC. obov Co + Carolin 14090 syp Calandrinia ( Baitaria) sect. Hirsutae Hook. & Arn. C. sericea Hook. & Arn. obov Co — Carolin 14121 syp C. umbellata (Ruiz & Pavon) DC. obov Sm — Boelcke 13859 syp C. uspalatensis Phil. obov Co - Carolin 14090 Syp PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 SYEDA SALEHA TAHIR AND ROGER CAROLIN 313 TABLE 1 (continued) Species name shape pattern aril Voucher Calandrinia (Baitaria) sect. Condensatae C. capitata Hook. & Arn. ob/or Sm — Carolin 14011 syp C. demissa Phil. obov Pp — CONC 8568 C. floribunda Phil. obov Co — Carolin 14109 syp C. glomerata Phil. ob/or Co — CONC 41891 C. leucocephala Phil. ob/or Co — Carolin 14108 syD C. modesta Phil. obov Co — CONC 32801 C. polycarpoides Phil. obov Co =") CONC 32732 C. prostrata Phil. obov Co — CONC 32747 C. trifida Hook. & Arn. ob/or Co/Sm — Carolin 14024 syp Calandrinia (Baitaria) sect. Acaules C. acaulis (Ruiz & Pavon) HBK. obov Pp + LIL 408812 C. affinis Gill. ex Arn. obov Sm/Co + Carolin 14013 syp C. affinis Gill. ex. Arn. obov Pp + Carolin 14014 syp C. caespitosa Gill. ex Arn. obov Co/Sm — Carolin 14122 syp C. fuegiana Gand. obov Co — CONC 32743 C. megarrhiza Helmsl. obov Co — RSA 631344 Calandrinia (Rumicastrum) sect. Pseudodianthoideae C. arenicola Syeda n.ob/or Ap + TYPEBRI 108559 C. balonensis Lindl. n.ob/or Co + AD 96923308 C. brevipedata F. Muell. ob/or Sm + TYPEMEL 48539 C. calyptrata Hook. f. ob/or Sm + AD 97314277 C. composita (Nees) Benth. n.ob/or Co + TYPE MEL 48615 C. corrigioloides F. Muell. ex Benth. ob/or Sm + TYPE MEL 48628 C. creethiae Morrison obov Sm-Co + C.A. Gardner 7821 PERTH C. disperma J. M. Black nr.obov Co-Pp — P.G. Wilson 8414 (see text) PERTH C. eremaea Ewart n.ob/or Pp + AD 96913076 C. granulifera Benth. obov Co + AD96929529 C. liniflora Fenzl ob/or Sm + H.Demarz 2725 PERTH C. monosperma Syeda obov Sm =) SUYPEIPERDE C. pickeringit Gray obov Pp-p + BRI015191 C. pleiopetala F. Muell. obov Co + TYPE MEL 48743 C. polyandra Benth. n.ob/or Co + AD 96343036 C. pumila F. Muell. obov Sm-Co + TYPE MEL 48789 C. quadrivalvis F. Muell. obov Co + TYPE MEL 48799 C. remota J. M. Black ob/or Sm P = AD 96416002 C. reticulata Syeda obov Rt P TYPENT 43058 C. sphaerophylla J. M. Black n.ob/or Co + TYPEAD 97826032 C. stagnensis J. M. Black obov Sm-Co + TYPEAD 96846192 C. strophiolata F. Muell. obov Sm + TYPE MEL 48826 C. volubilis Benth. n.ob/or Pp + TYPE MEL 48860 Calandrinia (Rumuicastrum) sect. Tuberosae C. lehmannii Endl. as-C Tb — Ll Setter427 PERTH C. primulzflora Diels cu Vv — MEL 48774 C. schistorrhiza Morrison cu Tb — 2 EeeVA pln 2434 PERTH Calandrinia (Rumicastrum) sect. Basales C. gracilis Benth. obov Co + BRI053297 C. papillata Syeda n.ob/or Pp + ‘TYPE PERTH C. plevopetala F. Muell. obov Co + TYPE MEL 48743 C. porifera Syeda obov Co + TYPEPERTH PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 314 SEED TYPES OF CALANDRINIA TABLE 1 (concluded) Species name shape pattern aril Voucher C. ptychosperma F. Muell. obov Rb + W.E. Mulham W941 NSW C. spergularina F. Muell. obov Co + TYPEMEL 48819 C. uniflora F. Muell. ob/or Sm — AD 96919061 Schreiterta S. macrocarpa (Speg.) Carolin oblong Co/pap — LiL Schreiter 4857 Abbreviations used for seed shapes: as-cl = asymmetric-clavate; cu = curved; n.ob/or = oblong to orbicular with a notch at insertion of funicle; ob/or = oblong to orbicular, + symmetric at base; obov = obovoid, + asymmetric at base; nr. obov = narrow obovoid. Abbreviations used for surface patterns: Ac = aculeate; Ap = aligned papillate; Co = colliculate; Po = polypiform; Pp = papillate; Pp-p = papillate-punctate; Rb = ribbed; Rt = reticulate; Sm = smooth; Tb = tuberculate; V = verrucate. Schreiteria is coded differently to indicate that the surface and shapes here are not comparable with the others in the table. DISCUSSION This survey indicates that there is range of seed types and seed surface patterns in Calandrinia sens. lat., and that they have a significant role to play in species recognition. Whilst the transformation series of seed shape and surface pattern are fairly clear, the polarization of these states, in the latter case at least, is not clear. The cladogram presented by Carolin (1987) indicates that the outgroup of the clade containing all of the segregate genera except Schreiterra is the Portulaca group of genera. This outgroup is equivocal for these characters. The outgroup of the whole family is Aizoaceae and little is published about the seed shape and surface patterns of this group. Shape There seems little doubt that curved shapes and those containing reduced endo- sperm are advanced features since they do not seem to occur in the Aizoaceae. The curved shape, together with the curved embryo and reduced perisperm, also occurs in the Portulaca group of genera (Carolin, 1987). The occurrence in the Portulaca group 1s considered to be a separate origin of the feature. We suggest that the oblong/orbicular and notched-oblong/orbicular shapes may represent the primitive condition whilst obovoid represents an asymmetry of the embryo at the base of the seed; the asymmetric-clavate and curved shapes represent an increas- ing asymmetry of the embryo. The narrow-obovoid shape is a special case found together with a narrow clavate shape in the heterospermic species, C. disperma, whilst Schreiterta macrocarpa, has an oblong seed with a reduced perisperm thus making the seed much narrower than the other oblong types. These are derivative forms of the obovoid or oblong shapes and are not equivalent to the curved type with reduced perisperm, since the embryo still almost encircles the perisperm. Seed surface pattern The cladogram presented by Carolin (1987) is equivocal with regard to the primi- tive state of seed surface pattern. Both papillate and smooth patterns occur in Calan- drinia (Baitaria) and evidence is lacking for Aizoaceae. Although it might seem more likely that the smooth condition is more primitive from a morphogenetic point of view, the evidence is slim. For the purposes of this description the smooth pattern 1s taken as a PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 SYEDA SALEHA TAHIR AND ROGER CAROLIN 315 starting point but some amendment of this will probably be necessary as more charac- ters of this group are considered. The colliculate pattern is a derivation of smooth by the development of the surface into a dome. Papillate represents an upgrowth from this dome with aculeate, tuber- culate and polypiform as further developments of that upgrowth. The papillate- punctate is a development from the papillate type as indicated above. The ribbed pattern is a development from the aligned papillate pattern as indicated above. The reticulate pattern is a quite separate development from colliculate/smooth patterns. Fig. 4 illustrates the putative evolutionary sequence in this character. VERRUCATE TUBERCULATE SMOOTH sees COLLICULATE == Pir PAPILLATE- a PUNCTATE ek PAPILLATE ACULEATE RETICULATE RIBBED POLYPIFORM Fig. 4. The suggested transformation series of seed surface pattern. Aril It is equally difficult to conclude whether the presence or absence of an aril is primi- tive since the outgroups of both the family (see Corner, 1976) and the clade containing Calandrinia sens. lat. (Carolin, 1987), have arillate as well as exarillate seeds. It is prob- able that the aril is a response to particular dispersal requirements and if this is so the lack of it is hkely to be primitive. The segregate genera of Calandrinia sens. lat. are consistent neither in seed shape nor surface pattern and neither of these attributes thoroughly characterizes them. Schreiteria is monotypic with a more or less unique seed type having a very curved embryo with very little perisperm and a mostly colliculate surface with a few blunt papillae. Calandrinia (Baitaria) consists mostly of species with colliculate or smooth surface patterns. Indeed, all those examined in sects Dianthoideae and Hirsutae have one of these two closely related patterns. One species in sect. Condensatae and two in sect. Acaules have papillate seeds. In fact, C. affinis at Farellones east of Santiago in Chile shows both papillate and colliculate surfaces in the same population. Calandrinia (Cistanthe) shows several surface types. Within sect. Cistanthe there is a definite trend from smooth/colliculate through papillate to aculeate and polypiform sur- face. Most of the species have one or other of the two most advanced states. This trend is not shown in any of the other segregate genera. Even sect. Amaranthoideae has only smooth-colliculate and papillate surfaces. Calandrinia sens. strict. shows both papillate and colliculate surfaces. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 316 SEED TYPES OF CALANDRINIA Within Calandrinia (Rumicastrum) it is doubtful if the sections of von Poellnitz (1934) are natural. Syeda (1980) has suggested that there are only three recognizable sections within Calandrinza in Australia but this also may be open to question. The only section which is more or less consistent is sect. ‘Iuberosae with a straighter embryo, less perisperm and a tuberculate or verrucate surface. Otherwise there would seem to be considerable homoplasy in these transformation series. The aril may be of more use at the sectional level. All the species of Calandrinia (Cistanthe) sect. Cistanthe examined so far have arils. Likewise Calandrinia (Cistanthe) sect. Amaranthoideae, Calandrinia (Baitaria) sects. Dianthoideae, Hirsutae and Conden- satae and Schreiterta do not have arils. The other taxa either have both conditions and/or have the indeterminate condition. It is clear, however, that one of the key characters provided by Pax and Hoffman (1935) to separate Yalinum and Calandrinia (sens. lat.), viz., the presence or absence of an aril, is not satisfactory. It seems, then, that the seed features are of limited value in distinguishing the segregate genera, although some genera and sections show distinct trends which, in the cases of Calandrinia (Cistanthe) and Calandrinia (Rumicastrum) sect. Tuberosae, are not repeated in other taxa at the same level. C. pickeringi and C. reticulata have unique surface patterns. The shape of the seed is also inconsistent within the segregate genera and sections except for Calandrinia (Rumuicastrum) sect. Tuberosae, Calandrinia (Cistanthe) sect. Cistanthe and Calandrinia (Baitaria) sect. Acaules. Schreiterta occupies an isolated position within Calandrinia sens. lat. Indeed Carolin (1987) has suggested it is more closely related to the Zaliznum group of genera although the seed characters neither confirm nor contradict this. It is clear from these results that the most variable of the segregate genera, with regard to seed shape and surface pattern, is Calandrinia (Rumicastrum). This greater variability is also shown in other characters such as fruit type and form of inflorescence. ACKNOWLEDGEMENTS We thank the Directors of the various Herbaria in Australia, Chile, Argentina, and U.S.A. who have lent material to us for this study from which seed samples have been taken. We thank also members of Universidad de Chile; Universidad de Concepcion; CEFAPRIN, Buenos Aires; Instituto Darwinion; Universidad de Cordoba and Instit- uto Lilloa for their help whilst RCC was engaged in collecting material for this study. In addition, we thank the staff of the Electron Microscope Unit at Sydney University for the use of their facilities, Tahir Rajput, Surrey Jacobs and Judy West for discussions on the work and Belinda Pellow and Cathy Miller for technical assistance. The University of Sydney provided funds which assisted in the investigation. References BLACK, J. M., 1948., — Flora of South Australia, ed. 2., vol. 2. Adelaide: Government Printer. CAROLIN, R. C., 1987. — A review of the family Portulacaceae. Austr J. Bot. 38: 383-412. , 1988. — Portulacaceae. Jn K. KUBITZKI1, (ed.), Families and Genera of Flowering Plants. In press. CORNER, E. J. H., 1976. — The Seeds of Dicotyledons, 2 vols. Cambridge: Cambridge University Press. CULLEN, D. C. ANON SUAREZ DE, 1953. — Las especies argentinas del genero Calandrinia (Portulacaceae). Bol. Soc. Arg. Bot. 5: 1-112. PAx, F., and HOFFMAN, K., 1935. — Portulacaceae. Jn A. ENGLER and H. HARMS, (eds), Die naturlichen pflan- zenfamilien, ed. 2, 16c: 233-262. Leipzig: Engelmann. POELLNITZ, K. VON., 1934. — Die Calandrinien-Arten Australiens. Fedde Rep. 35: 161-173. REICHE, K., 1888. — Flora de Chile, vol. 11. Santiago: Universidad de Chile. STEARN, W. T., 1966 — Botanical Latin. London: Nelson & Sons. SyeEDA, S. T., 1980 — The genus Calandrinia HBK. in Australia. Sydney: University of Sydney, M.Sc. thesis, unpubl. PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 A late Holocene Vegetation and Fire Record from Ku-ring-gai Chase National Park, New South Wales P. G. KODELA and J. R. DODSON Kope_a, P. G., & DopDson, J. R. A late Holocene vegetation and fire record from Ku- ring-gai Chase National Park, New South Wales. Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 317-326. Pollen and charcoal analyses of sediments from South Salvation Creek Swamp in Ku-ring-gai Chase National Park indicate that pollen influx has been dominated by local swamp species and dry sclerophyll heath and woodland taxa for the last 6000 radiocarbon years. Fire occurred throughout the record but charcoal and Eucalyptus pollen influx decreased over the last 1700 years. In an environment supporting dry sclerophyll vegetation fire appeared to play a constant role. The swamp surface was initially a sedgeland but was invaded by Glezchenia and woody shrubs around 2500 b.p. None of the vegetation changes could be ascribed directly to climatic shifts but the origin of the swamp itself may have been due to the postglacial rise in sea-level. P.G. Kodela and J. R. Dodson, School of Geography, University of New South Wales, PO. Box 1, Kensington, Australia 2035, manuscript received 19 January 1988, accepted for publication 20 April 1988. INTRODUCTION Ku-ring-gai Chase National Park encompasses 14,700ha of natural bushland 20km north of Sydney. A palynological investigation was undertaken on a peat swamp deposit on the southern arm of Salvation Creek on Lambert Peninsula (Fig. 1) to investigate vegetation change and possible vegetation-fire relationships during the mid to late Holocene. The elevation of the site (130m a.s.1.) assures that it has remained beyond any direct effects of sealevel change. Ku-ring-gai Chase experiences a temperate coastal climate with a mean annual rainfall exceeding 1200mm. Mean maximum temperatures range from 26°C in Janu- ary to 16°C in July whilst mean minima range from 17°C in January to 7°C in July. The prevailing winds are northeast to northwest in summer and southwest to southeast in winter (Bureau of Meteorology, 1979; Fitzpatrick and Armstrong, 1972; National Parks and Wildlife Service, 1982). The rugged topography of Ku-ring-gai Chase is strongly influenced by the predominant rock type, Hawkesbury Sandstone. Shales and sandstones of the under- lying Narrabeen Group outcrop along the foreshores of the Hawkesbury River. Lambert Peninsula is characterized by deeply dissected V-shaped valleys with narrow divides on the west and extensive areas of low slope on the plateau surface and steep coastal gullies on the east (Buchanan, 1975, 1980). Pittwater forms part of the drowned river valley of the lower Hawkesbury River. The study site, South Salvation Creek Swamp, has developed in a broad shallow valley on the gently sloping plateau. Drainage has been impeded by massive quartz sandstone and clayey layers derived from shale lenses within the Hawkesbury Sandstone group. Lamy and Junor (1965a) described the importance of such swamps in maintain- ing a perennial flow to creeks, reducing flood flows on the lower sections, controlling erosion, and arresting sedimentation. Soils derived from siliceous Hawkesbury Sandstone tend to be light-coloured sandy loams with low humus content, poor water-retaining properties and low fertility PROC. LINN. SOC. N.S.W,, 110 (4), (1988) 1989 318 A LATE HOLOCENE VEGETATION AND FIRE RECORD 151°15' ie BROKEN BAY Eleanor Bluffs 35° a Challenger x Head «Taylors Point LEGEND 222 Salvation Creek catchment £9 South Salvation Creek swamp £3 Urban or cleared areas 0 1 2 kilometres 151°15' (| ‘oor Fig. 1. Location of South Salvation Creek Swamp showing core location. (Pidgeon, 1937; Lamy and Junor, 1965a, 1965b). On steep slopes and rocky outcrops sandy skeletal soils (lithosols) occur while deeper soils with some degree of profile development (towards yellow podzolics) are associated with the more gentle plateau slopes and hillslopes. Fine-textured loams of relatively higher-nutrient status, water- retaining capacity and humus content have developed on the Narrabeen shales. The nature and distribution of the vegetation are strongly influenced by geology, topography, soil, drainage, aspect and fire (Buchanan, 1975, 1980; Kodela, 1984; Outhred ¢ al., 1985; Thomas and Benson, 1985). The dissected terrain and changes in lithology result in a complex mosaic of habitats and thus plant communities vary markedly in structure and floristics. The Salvation Creek catchment is dominated by woodland and heath on the exposed plateau surface and upper hillslopes while taller more mesic forests are confined to the sheltered lower slopes and gullies fronting Pittwater. Descriptions are based on Thomas and Benson (1985) for terrestrial communities and Buchanan (1980) and Kodela (1984) for other types. Closed-scrub and heath occur in poorly-drained areas on the plateau, and shallow sandy soils on rocky sandstone out- crops. Shrub species commonly associated with moist heath include Banksia ertcifolia, PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. G. KODELA AND J. R. DODSON 319 Allocasuarina distyla and Hakea teretifolia. Occasionally Eucalyptus haemastoma and Eucalyp- tus gummifera occur as emergents. Low woodland and low open-woodland occur on the better drained lithosols and sandy clay loams on ridges, crests and low to moderately sloping benched hillslopes. Common tree species include E. gummifera, E. haemastoma and E. oblonga often with a diverse shrub layer characterized by species from the families Epacridaceae, Fabaceae, Myrtaceae, Proteaceae and Rutaceae. Sheltered slopes with south to southeast aspects support open-forest of E. piperita ssp. piperita, Angophora costata and E. gummifera. The deeper more fertile soils on the shale and colluvium-derived soils of the lower slopes support Eucalyptus maculata, E. paniculata, Allocasuarina torulosa and Syn- carpia glomulifera. Sheltered coastal gullies may contain small pockets of rainforest with Ceratopetalum apetalum, Tristaniopsis laurina, Acmena smithi, Livistona australis, Synoum glandulosum and Elaeocarpus reticulatus. South Salvation Creek Swamp has a complex of four swamp vegetation types (Buchanan, 1980). The dense cover is essentially sedge, fern and shrub-dominated. Below the tall (2-3m) emergent Leptospermum juniperinum there is a layer of Banksza robur, Aotus ericordes and Gahnia sieberiana. The surface layer is dominated by Empodisma minus, Gleichenia dicarpa and Drosera binata. Other swamp and moist heath species, often peripheral to the swamp, include Gymnoschoenus sphaerocephalus, Xyris operculata, Lepidosperma sp., Sprengelia incarnata, Hakea teretifolia and Banksia errcifolia. METHODS A vertical core of peat was extracted from the deepest sediment section with a Russian (D-section) corer (Jowsey, 1966). Subsamples at about 10cm intervals were dis- persed in warm hydroxide, treated with hydrofluoric acid and acetolysis fluid (9 acetic anhydride: 1 concentrated sulphuric acid) and dehydrated through an alcohol series before the residue was mounted in silicone oil. The technique is based on that described by Erdtman (1969), Faegri and Iversen (1975) and Moore and Webb (1978). A minimum of 200 pollen grains and spores was counted for each sample depth and pollen sums for the percentage diagrams are based on the total count. Five peat samples were radiocarbon-dated to provide dates for the age-depth curve in Fig. 2 and the time scale in Figs 3 and 4. Loss-on-ignition was used as an estimate of organic content, and, using the point count method of Clark (1982), microscopic charcoal particles were counted as an index of fire activity. RESULTS AND DISCUSSION Stratigraphy The core was predominantly herbaceous peat overlying a pallid inorganic layer of sand and clay. Within the peat the degree of humicity increased with increasing depth, from course fibrous peat at the surface to well-humified organic matter at the base. The gradual transition suggests a series of decay. The sediment can be summarized as follows: Depth (cm) Description 0-20 dense surface mat of undecomposed plant remains (mainly Empodisma and Glezchenia): 20-40 mainly water and living roots. Gradual transition to: 40-110 brown fibrous peat ( Turfa herbacea), pH 4.5-5. Gradual transition to: 110-240 homogeneous fine fibrous clayey peat with charcoal fragments and quartz sand scattered throughout or in thin layers; pronounced H,S odour. Distinct granular charcoal layer at base of peat (230-240cm). Sharp transition to: 240+ grey light-medium sandy clay (Grana/Argilla). Bedrock was encoun- tered at ca 285cm. PROG. LINN. SOC. N.S.W., 110 (4), (1988) 1989 320 A LATE HOLOCENE VEGETATION AND FIRE RECORD Age and Origin of Site The organic sediment dates from between 5000 and 6000 b.p. and marks either a regional change to greater effective precipitation, probably as a result of higher post- glacial sealevel, or a local geomorphic event such as damming of a stream outlet to allow sedimentation to occur. There is no clear evidence of the latter however. Buchanan (1975, 1980) suggested that the swamps of the Sydney region may be representatives of a cycle of swamp build-up and destruction. She suggested that climatic changes appear to be responsible for the cycles; and even in the short term such fluctuations over decades can affect swamp boundaries. Very dry climatic conditions can crack the peat surface, enable fires to burn the sediments and promote erosion when followed by heavy rain. The charcoal layer at the base of the peat (1.e. peat-inorganic boundary) may be an in- dication of such a process; alternatively the charcoal may have been stored in the catch- ment until deposition and preservation began. Such substantial layers of charcoal at the base of other peat or clay deposits have been observed by several investigators (Clark and Colhoun, pers. comm.; Macphail, 1984; Dodson et al., 1986) and may be due to flush of charcoal into the system once sediments are able to accumulate or indeed may be part of a changed environment which leads to sedimentation itself. On a smaller scale, possibly reflecting periods of drought and fire, scoured shallow depressions are common on the exposed Hawkesbury Sandstone plateau while other depressions are inhabited by mosses, herbs and small shrubs which build up soil and organic matter. Due to the general lack of distinct layers within the peat radiocarbon dates were not related to stratigraphy. Fig. 2 shows a general increase in accumulation rates toward the present, due at least in part to compaction increasing with depth. It is from this curve that ages in the text are estimated. Pollen and Charcoal Record Fig. 3 shows abundance of pollen for individual taxa based on percentage of the total pollen sum for each sampled sediment depth; in Fig. 4 estimates of pollen influx are shown for the more abundant pollen types. In these diagrams the modern sample incor- porates the uppermost 15cm of sediment. Samples at 40 and 45cm represent 5cm sample thicknesses, 1.e. 40-45cm and 45-50cm respectively, while all other samples represent lcm sample thicknesses at 10cm intervals. Influx values are not calculated for the surface or more minerogenic samples; and there are missing loss-on-ignition values at several depths. The Appendix shows likely genera and species represented by the pollen taxa on the diagrams. Tree pollen was dominated by Eucalyptus, but the values for Allocasuarina and Banksia may also contain some representation from trees. The highest values for Eucalyptus occur prior to 1900 b.p. for both influx and percentage values. Charcoal is present throughout the record with highest values for both influx and concentrations between about 1300 b.p. and 3000 b.p. Charcoal concentration shows a decline from about 500 b.p. There is no consistent small-scale correlation between Eucalyptus and charcoal but the highest values for both occur prior to 1900 b.p. Dryland herbs are poorly represented. These include Asteraceae, Gonocarpus, Lilia- ceae, Poaceae and Pteridium. An increase in herb pollen influx after 2000 b.p. may represent disturbance as many species in this group are competitive when clearance occurs. Although some of the herb taxa show slightly higher abundance values cor- responding to relatively high fire activity, as indicated by charcoal, in this part of the record there are no consistent associations. The total pollen input into the site is dominated by swamp taxa such as Glezchemza, Cyperaceae and Restionaceae; and is therefore very local in nature. There is a general trend of increasing Gleichenia spores to the present while Cyperaceae may have PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P.G. KODELA AND J. R. DODSON ai Age - “Radiocarbon years B.P. 0 1000 2000 3000 4000 5000 50 + 0.16 cm yr7! 100 + 0.04 cm yr~' + 0.06 cm yr-'! 200 + 0.02 cm yr! 250 * + Two standard deviations + mean accumulation rates Fig. 2. Age-depth curve for the peat sediments. decreased slightly. Restionaceae pollen increases markedly at 4900 b.p. where the peat layer overlies the inorganic sandy clay. Influx values for Glezchenia, Cyperaceae and Restionaceae appear somewhat cyclic, generally following changes in charcoal influx. Influx peaks for these taxa around 1700 b.p. and 500 b.p. correspond to peaks in char- coal and may indicate fires on or near the swamp promoting growth of these taxa. Past Vegetation and Environmental History There have been no marked changes in the flora; all the major pollen taxa remained PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 322 A LATE HOLOCENE VEGETATION AND FIRE RECORD KU-RING—GAI CHASE NATIONAL PARK: SOUTH SALVATION SWAMP Pollen Sum: total pollen count Age Depth BP. (cm) 0 10% Age b.p. Modern 0 | 1067 50 Sales 1000 il cp 2000 25404 9004 anag 51204 5000 ae Eucalyptus Monotoca Exocarpus Hibbertia Leptospermum/Baeckea Other Epacridaceae Sauera Other Myrtaceae Fabaceae type Myrsinaceae Gyrostemonaceae Allocasuarina Acacia Rutaceae A Depth Banksia Podocarpus Pomaderris San ce Other Proteaceae Dodonaea Coprosma B.P. (cm) Age bp. Modern 7 0 | 106 | 50 p E pass eral nog 1000 Ae ee f : t ! 2000 + { sie | | 8 51204 j 5000 | 2504 | E ’ Brassicaceae Scrophulariaceae Asteraceae (Lig.) Myriophyllum Euphorbiaceae Loranthaceae Asteraceae (Tub.) Drosera Tetratheca Chenopodiaceae F/antago varia Cyperaceae Mitrosacme Polygalaceae Poaceae Restionaceae A Geraniaceae Apiaceae Liliaceae ge Depth Malvaceae Ranunculaceae Gonocarpus B-P. cm) Acelorp! Modern 7 0 lien 106 | 504 ie i oe 1000 650 ‘al = f | 2000 4 5120 j 5000 zat | | Gleichenia Flantago lanceolata Fern spores Tricolporate grains Pteridium Charcoal concentration («10 000 particles/cm%) Pinus 7 Loss on Ignition Echium Aumex Fig. 3. Percentage pollen diagram from South Salvation Creek Swamp. In this diagram the modern sample incorporates the uppermost 15cm of sediment. Samples at 40 and 45cm represent 5cm sample thicknesses, ie. t0-45em and 45-50cm respectively, while all other samples represent lem sample thicknesses at 10cm intervals. present throughout the record. The sandstone flora has retained its sclerophyll nature, reflecting stability and therefore resilience to nutrient-poor sandy soils, high insolation PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. G. KODELA AND J. R. DODSON 323 KU-RING-GAI CHASE NATIONAL PARK: SOUTH SALVATION SWAMP Influx diagram A Depth BP. (em) 9 100 gt/em2/yr Age bp. Modern 4 0) r 1064 50 ae ES | 1000 1650 | 150 lead 2540 3000 200 t 4000 21204) 204 Eucalyptus Leptospermum/Gaeckeo Other Myrtaceae Banksia Allocasuarina Other Proteaceae Monotoca Other Epacridaceae Age Depth BP. (cm) Age b.p Modern > 0 1064 50 415 ao 1000 1650 150 2000 25404 3000 200 7 4000 5120 5000 250 Fabaceae type Cyperaceae Restionaceae lodonaea Asteraceae (Tub.) Poaceae Liliaceae Be. (Gn) Conocarpus Age bp. Modern 0 10647 50 4154 | 100 1000 1650 fd 2000 25404 3000 700 4000 51204 5000 250 Gleichenia Fern spores Pteridium Charcoal influx (*10 particles /cm2/yr) Fig. 4. Influx diagram for major pollen types from South Salvation Creek Swamp. exposure, drought, and fire. Such vegetation is probably unlikely to be sensitive to minor environmental changes. Although the results show few or no changes in vegeta- tion composition there are changes in relative abundances. The pollen of many of the heath and understorey shrubs of woodland and forest communities is poorly dispersed, including that for Acacia, Epacridaceae, Fabaceae, Proteaceae and Rutaceae (Dodson, 1983). This explains the relatively low values for these taxa which are common in the vegetation at Ku-ring-gai Chase. Their under- representation in the pollen record does not reflect their true importance in the vegetation. Fires appear to have been present throughout the record, reflecting a consistent association between fire and dry sclerophyll vegetation. There is a decline in charcoal within the last 200 years, coinciding with the arrival and settlement of European Man in the Sydney region. This may reflect the displacement of Aborigines and their burning practices. At the same time European presence is marked in the surface samples by Echium, Pinus, Plantago lanceolata and Rumex pollen. Low loss-on-ignition values corresponding to thin sandy lenses appear with or above (indicating possible time lags) relatively high charcoal levels at around 4200 b.p. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 324 A LATE HOLOCENE VEGETATION AND FIRE RECORD and 2300 b.p. These are likely to indicate disturbance and erosion from the surrounding slopes. Trends in pollen of swamp taxa show that the early swamp surface was dominated by Restionaceae, and possibly a fern other than Glezchenia during the more minerogenic deposition. Woody shrubs: Banksia robur, Leptospermum juniperinum and Aotus ericordes, prefer deeper sediments which could explain their general increases toward the present as a local successional effect. Increasing deposition would have created drier conditions on the swamp favouring their establishment and that of Gleichenia. Above this general trend smaller changes probably reflect local variations in water-table depth or the changing patterns of distribution of taxa by population fluctuations, interspecific competition, or fire. Any relationship between charcoal influx and pollen abundance from swamp taxa could reflect fire adaptations of these taxa. Although normally much regeneration would be from rhizomes and suckering, the clearance of the thick Gleichenia and Empodisma cover would allow initial recruitment by seed dispersal, particularly of woody shrubs. Nutrient influx to the swamp after fires may also have some impact on the swamp vegetation. Fire may therefore be an important factor in swamp vegetation dynamics. A peak in Leptospermum pollen occurs with a decline in Cyperaceae, Restion- aceae and Gleichenia around 2900 b.p. which could represent a dry period allowing L. juniperinum to expand on the swamp. The initial major decline in Eucalyptus around 1800-1900 b.p. corresponds with a trend of increasing Glezchenia and swamp shrubs which may together represent climatic conditions becoming slightly drier and the eucalypt canopy opening towards a more woodland type. Alternatively the mire surface could have expanded laterally, thereby pushing back the eucalypt woodland and reducing the local eucalypt pollen input reach- ing the core site. Eucalyptus values may have been high in the past due to trees directly overhanging the site. Chalson (1983) studied the vegetation record of Jibbon Swamp in Royal National Park south of Sydney. The site is located on a sandsheet overlying Hawkesbury Sand- stone with the swamp set in the swale of a U-shaped dune about 1km from the shoreline and 10m above sea level. Chalson (pers. comm.) found Eucalyptus to be more common prior to about 5000 b.p. with E. szebert pollen (possibly the mallee form of E. szebert which occurs in the area today) highest around 4000 b.p. and declining to the present. At Kur- nell Peninsula Eucalyptus declined after 2000 b.p. and was replaced by dune shrubs such as Leptospermum and Monotoca (Martin, 1986, and 1988 pers. comm.). These studies may suggest a regional decline in Eucalyptus in the late Holocene, a question worthy of further research. The results show a very high local pollen input from plants growing on or near the swamp. For densely vegetated sites like South Salvation Creek Swamp it is difficult to separate terrestrial from swamp pollen where several genera have species representative in both habitats. For example, the source of Leptospermum pollen could be from the many woodland and heath species surrounding the swamp, however most of the pollen in the record is likely to be derived from Leptospermum juniperinum growing on the swamp. Such results emphasize the problems encountered when interpreting a pollen record derived from a complex mosaic of vegetation communities surrounding a swamp. The lack of any consistent relationship between charcoal abundance and the behaviour of individual taxa reflects the complex relationships between vegetation and fire and the input of pollen and charcoal in the sediment. It may well be that the catch- ment area for charcoal differs from that of the pollen. Plant taxa and communities are likely to have evolved and adapted to particular fire regimes rather than fire per se (Gill, 1975). Studying the effects of fire on vegetation PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P. G. KODELA AND J. R. DODSON 325 requires separating the components of the fire regime (intensity, frequency, type, time of occurrence etc.) and examining these in relation to the adaptations and life cycles of the plants (Kodela, 1984). Palynological techniques cannot readily differentiate these. Nevertheless broad relationships can be established and in this case there appears to be at least an association between high Eucalyptus and charcoal influx; perhaps reflecting an interdependence between the eucalypt species at the site and fire. Although sclerophyll and swamp taxa appear to have dominated the record, fluctu- ations in their abundances and/or distributions have occurred. These are likely to be results of a combination of factors, including water-table fluctuations, seasonal drought, fire activity, interspecific competition, the natural changing patterns in species distri- bution and the impact of Aboriginal and European people. Changes in swamp and ter- restrial pollen taxa around 2000 b.p. may indicate a drier climate to the present. ACKNOWLEDGEMENTS We are grateful to Chris Myers (UNSW) for technical help, Colin Pain (UNSW) and Bob Conroy (NPWS) for advice and field assistance, Kevin Maynard and John Owen (UNSW) for cartography and Lynne Illidge for typing the manuscript. Many friends from the 1984 honours class in Physical Geography group kindly provided labour in the field. The N.S.W. National Parks and Wildlife Service (NPWS) generously provided funds for radiocarbon dating. References BUCHANAN, R., 1975. — The Relationship of Swamp, Shrub and Podzol Distribution to the Geology and Physiography on the West Head Peninsula. North Ryde (N.S.W.): Macquarie Univ., B.Sc. honours thesis, unpubl. ——, 1980. — The Lambert Peninsula, Ku-ring-gai Chase National Park. Physiography and the distri- bution of podzols, shrublands and swamps, with details of the swamp vegetation and sediments. Proc. Linn. Soc. N.S.W. 104: 73-94. BUREAU OF METEOROLOGY, 1979. — Climatic Survey Sydney, Region 5, New South Wales. Canberra: Department of Science and the Environment. CHALSON, J., 1983. — Palynology and Paleoecology of Jibbon Swamp, Royal National Park, N.S.W. Kensington (N.S.W.): Univ. of N.S.W., B.Sc. honours thesis, unpubl. CLARK, R. L., 1982. — Point count estimation of charcoal in pollen preparations and thin sections of sedi- ments. Pollen et Spores 24: 523-535. Dopson, J. R., 1983. — Modern pollen rain in southeastern New South Wales, Australia. Rev. Palaeobot. Falynol. 38: 249-268. ——, GREENWOOD, P. W., and JONES, R. L., 1986. — Holocene forest and wetland dynamics at Barrington Tops, New South Wales. Journal of Biogeography 13: 561-585. ERDTMAN, G., 1969. — Handbook of Palynology. Copenhagen: Munksgaard. FAEGRI, K., and IVERSEN, J., 1975. — Textbook of Pollen Analysis. Copenhagen: Munksgaard. FITZPATRICK, E. A., and ARMSTRONG, J., 1972. — The bioclimatic setting. Proc. Ecol. Soc. Aust. 7: 7-26. GILL, A. M., 1975. — Fire and the Australian flora: a review. Aust. For., 38: 4-25. Jowsey, P. C., 1966. — An improved peat sampler. New Phytol. 65: 245-249. KODELA, P. G., 1984. — The Vegetation and Fire History of Ku-ring-gai Chase National Park. Kensington (N.S.W.): Univ. of N.S.W., B.Sc. honours thesis, unpubl. Lamy, D. L., and JUNoR, R.S., 1965a. — An erosion survey in the Ku-ring-gai Chase and adjoining catch- ments. II. J. Sor! Cons. N.S.W. 21: 159-174. , and , 1965b. — An erosion survey in the Ku-ring-gai Chase and adjoining catchments. I. Survey of the Cowan Water and Cockle Creek catchments. J. Soil Cons. N.S.W. 21: 94-110. Macpual_L, M. K., 1984. — Small-scale dynamics in an early Holocene wet sclerophyll forest in Tasmania. New Phytologist 96: 131-147. Martin, A. R. H., 1986. — Palaeoecology, palaeolimnology, palynology and coastal environment-areas of neglect in coastal studies? Jn: E. FRANKEL, J. B. KEENE, and A. E. WALTHO, (eds), Recent sediments in eastern Australia: marine through terrestrial. Sydney: Geol. Soc. Aust., NSW Division, Publ. No. 2. Moore, P. D., and WEB, J. A., 1978. — An Illustrated Guide to Pollen Analysis. London: Hodder and Stoughton. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 326 A LATE HOLOCENE VEGETATION AND FIRE RECORD NATIONAL PARKS AND WILDLIFE SERVICE, 1982. — Teachers Handbook to Ku-ring-gai Chase National Park. Sydney: National Parks and Wildlife Service of N.S.W. OUTHRED, R., LAINSON, R., LAMB, R., and OUTHRED, D., 1985. — A floristic survey of Ku-ring-gai Chase National Park. Cunninghamia 1: 313-338. PIDGEON, I. M., 1937. — The ecology of the central coastal area of New South Wales. I. The environment and general features of the vegetation. Proc. Linn. Soc. N.S.W. 62: 315-340. THOMAS, J., and BENSON, D. H., 1985. — Vegetation Survey of Ku-ring-gai Chase National Park. Sydney: National Herbarium of New South Wales. APPENDIX THE LIKELY ORIGINS OF THE MAIN TAXA IN THE POLLEN DIAGRAMS Taxa identified in the pollen and spore record Eucalyptus (includes Angophora spp.) Leptospermum/Baeckea Myrtaceae (other) Allocasuarina Banksia Proteaceae (other) Monotoca Epacridaceae (other) Fabaceae Euphorbiaceae Liliaceae Gonocarpus Cyperaceae Restionaceae Gleichenia Pteridium Fern spores Pinus The main or most likely genera and species represented by the preserved pollen and spores, and their ecology Eucalyptus haemastoma, E. gummifera, E. oblonga, E. punctata, E. umbra, Angophora costata etc. Forest and woodland trees. Would include Leptospermum juniperinum which grows on the swamp, L. squarrosum which occurs in moist heath communities and Baeckea spp., etc which occur in heath communities or as understorey shrubs in woodland and forest communities. Shrubs including Kunzea, Callistemon and Melaleuca. A. distyla (tall shrub to small tree in moist or dry heath communities), A. torulosa, and A. /ittoralis (trees in sheltered forest and woodland communities). Banksia robur (shrub growing on the swamp), B. serrata, B. marginata and B. spinulosa (common shrubs in woodland and heath communities), B. ericifolia (moist heath shrub). Hakea teretifolia (moist heath shrub) Jsopogon anethifolius, Petrophile pulchella, Conosper- mum spp., Grevillea spp. etc. Shrubs in the forest, woodland and heath formations. Monotoca elliptica, M. scoparia. Shrubs in woodlands and heath. Some species occur on the swamp margin e.g. Sprengelia incarnata. Leucopogon (low heath or woodland shrub) encountered several times in the pollen record. Aotus ericoides (woody shrub growing on the swamp) and many heath and woodland species such as Dillwynia spp., Pultenaea spp., Bossiaea spp., Phyllota phylicoides. Mainly Amperea xiphoclada (low shrub) Xanthorrhoea spp., Thysanotus spp., Dianella spp. Herbs in heath, woodland and forest communities. G. tetragynus, G. teucrioides, G. micranthus. Herbs in heaths and forests. Gahnia sieberiana, Lepidosperma spp., Gymnoschoenus sphaerocephalus, Caustis spp. Swamp and moist heath sedges. Empodisma minus, Restio spp. Swamp herbs. (Also on moist rock shelves and in moist heaths). Gleichenia dicarpa. Fern growing on swamp and in other moist areas. Pteridium esculentum. Often increases with disturbance. These include Cyathea, Lycopodium, Blechnum, Polypodiaceae, Adiantum and Selaginella. Understorey species in moist forests, near creeks, moist rock crevices, etc. Pinus radiata. Introduced trees. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 A new Species of Melita (Crustacea: Amphipoda: Melitidae) from northern New South Wales with a Note on the Genus Abludomelita Karaman, 1981 W. ZEIDLER (Communicated by F. W. E. Rowe) ZEIDLER, W. A new species of Melita (Crustacea: Amphipoda: Melitidae) from north- ern New South Wales with a note on the genus Abludomelita Karaman, 1981. Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 327-338. A new species of Melita from a small, artificial, freshwater coastal lake at Angourie, northern N.S.W. is described. It was found with other fauna characteristic of the larger coastal rivers of eastern Australia. It is most similar to the New Zealand species M. awa Barnard, 1972, but does not share the same pleonal tooth formula. M. festiva (Chilton 1885), the only Australian species with the same pleonal tooth formula, differs pri- marily in the form of the gnathopods and the small posteroventral tooth of the third pleonal epimeron. The new species does not readily fit into either Melzta or Abludomelita as defined by Karaman (1981) and the division of Melita into the above two genera 1s considered unjustified without a more detailed revision. W. Zeidler, Marine Invertebrates, South Australian Museum, North Terrace, Adelaide, Australia 5000; manuscript received 18 February 1987, accepted for publication 20 April 1988. INTRODUCTION There has been no comprehensive study of the genus Melita in Australia and of the 61 species listed by Barnard and Barnard (1983) only four have been recorded from Aus- tralia. Sheard (1937) only listed M. festiva and M. fresnel (now in Dulichiella). Barnard (1972) described two new species, M. matilda and M. oba and a new subspecies of M. zeylanica (keuertz) all from south Western Australia but did not list any other species of Melita. M. zeylanica, a cosmopolitan species, has been recorded from Australia more recently by Collett et a/. (1981). Kangas and Geddes (1984), Poore (1982) and Potter et al. (1981). The species described here was found in a small, coastal, freshwater lake (actually a water-filled, disused quarry), known as “The Blue Pools, at Angourie, 5km south of Yamba, northern New South Wales. It was found near the shallow edges together with a small hydrobiid snail (Posticobia brazierx) and the hymenosomid crab (Amarinus lacustris) both of which are characteristic of the fauna of larger coastal rivers of eastern Australia. MATERIALS AND METHODS Specimens were collected from the shallow edges of only one of the two small fresh- water lakes at Angourie. The species was not particularly abundant and only 10 males and 5 females were collected. I was unable to obtain specimens from coastal rivers nearby and no additional material was available from the Australian Museum, Sydney. The holotype male and allotype female are deposited in the Australian Museum, Sydney (AM), and the remainder, all designated paratypes, are deposited in the South Australian Museum, Adelaide (SAM). Specimen length was measured along a lateral parabolic line drawn from the anterior extremity of the head through the middle of the body to the posterior limit of pleonite 6. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 328 A NEW SPECIES OF MELITA Y il, Fy yy Wh fp ADH A WIA) Fig. 1. Melita plumulosa sp. nov. Holotype male; scale bars = 1mm and 0.2mm respectively. Abbreviations used in figures and text: Al, antenna 1: A2, antenna 2; Mxp, maxilliped; Mx1, maxilla 1; Mx2, rnaxilla 2; Md, mandible; UL, upper lip; LL, lower lip; G1, gnathopod 1; G2, gnathopod 2; P3-P7, pereopod 3 to pereopod 7; U1-U3, uropod 1 to uropod 3; T, telson; P14, P15, fourth and fifth pleonite; R, right; L, left. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 W. ZEIDLER 329 Unless indicated otherwise dissected appendages were taken from the left hand side of the animal. All specimens and dissected appendages are preserved in 2% formalde- hyde/propylene glycol solution. Material examined Holotype 6 (AM, P37289), allotype 9 (AM, P37290) and 11 paratypes (96,4 9 ) (SAM, C4076), all from the shallow edges of the northern-most pond (freshwater) of ‘The Blue Pools’ at Angourie, 5 km south of Yamba, N.S.W., collected by W. Zeidler, 4.1X.1984. Also examined were all specimens of Melzta in the collections of AM and SAM. DESCRIPTION Melita plumulosa sp. nov. (Figs 1-5 ) DESCRIPTION OF HOLOTYPE: Male 6.7mm (AM, P37289). Head (Fig. 1): Rostrum absent, length slightly less than first two pereonites combined; lateral cephalic lobes evenly rounded; post-antennal sinus short; eyes black, ovato-circulate. Antenna 1 (Fig. 2): Almost as long as body; all articles with long irregularly plumulose setae particularly on posterior margin; article 1 about 80% length article 2, expanded near base with three ventral spines; article 3 short, about half length article 2; accessory flagellum 3-articulate plus tiny fourth article (2 + 1, L); flagellum with 18 + (R, broken), 17 (L) articles. Antenna 2 (Fig. 3): Little more than half length antenna | with similar but generally longer setae; gland cone enlarged reaching mid-way peduncular article 3; penduncular article 4 longest almost twice length article 5; flagellum with.9 articles (L & R) together about % length peduncular article 4. Upper Lip (Fig. 4 ): Broader than long, apically rounded, bearing numerous short setae distally. Lower Lip (Fig. 4): Inner lobes barely developed; outer lobes with medial tooth-like projection and fringe of short setae. Mandibles (Fig. 4): Palp 3-articulate, article 2 about twice length article 1 with one long plumulose seta medially, article 3 slightly shorter than 2 with three short and one very long plumulose seta terminally; incisor with five teeth; lacina mobilis with four teeth; spine row with four spines, the one nearest lacina mobilis spatulate; molar triturative with one long plumulose seta. Maxilla 1 (Fig. 4): Inner lobe with six apical plumulose setae; outer lobe with six apical trifurcate spines; palp 2-articulate with dilated, distal article armed with six terminal teeth and eight sub-terminal setae. Maxilla 2 (Fig. 4): Inner lobe shorter and narrower than outer with about twenty setae apically and marginally; outer lobe with about twelve strong apical setae. PROG. LINN. SOC. N.S.W., 110 (4), (1988) 1989 330 A NEW SPECIES OF MELITA Fig. 2. Melita plumulosa sp. nov. Holotype male; scale bar = 0.2mm. Sec legend to Fig. 1 for key to abbreviations PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 W. ZEIDLER 331 Maxilliped (Fig. 4): Inner lobe reaching % along article 2 of palp bearing several setae apically and along inner margin; outer lobe reaching 3/5 along article 2 of palp, semi- circular about half as wide again as inner lobe with seven apical feathered setae and two rows of shorter setae along inner margin interspersed with bristles; palp article 2 with plumulose setae along inner margin, article 3 about % length article 2 dilated distally with patch of close-set bristles on inner distal corner and four long setae on outer distal corner, article 4 unguiform, subequal to article 3 with terminal nail and fringe of short setae on inner margin. Gnathopod 1 (Fig. 1): Coxal gill absent; coxa with plumulose setae on lower and distal half of anterior margin; article 6 is 60% length article 5 with anterior hump, palm trans- verse with medial hump extending to outer margin of dactylus; dactylus curved, blunt with three strong setae on postero-distal corner, shorter than width of palm. Gnathopod 2 (Fig. 2): Less setose and much larger than gnathopod 1; coxal gill sac-like, slightly longer than wide, as long as article 2 with very convex posterior margin and almost straight anterior margin; coxa slightly larger but like that of gnathopod 1; article 4 with sharp posterodistal extension; article 6 unlobed, large, about % length articles 2- 5 combined, palm oblique with few scattered plumulose setae otherwise smooth and undefined by teeth or processes; dactylus smooth length equal to width article 6, over- riding palm on inside face, fitting into weak undefined hollow. Pereopod 3 (Fig. 2): Slightly shorter than gnathopod 2; coxal gill like that of gnathopod 2 but slightly longer and narrower; coxa slightly larger than that of gnathopod 2 with slightly concave posterior margin; articles 5 and 6 subequal in length slightly shorter than article 4; all articles sparsely setose. Pereopod 4 (Fig. 2): About % length pereopod 3 otherwise similar except for following; coxa more rectangular, wider and shorter; article 6 slightly longer than article 4 or 5 which are subequal in length. Pereopod 5 (Fig. 3): About twice length pereopod 4; coxal gill oval-shaped, smaller than that of pereopod 4, about % length article 2; coxa wider than article 2 with large anterior lobe equivalent to about 40% of total coxal length and small posterior lobe; article 2 rectangular about % as wide distally as proximally with conical posterodistal projection; article 6 about 1% times length article 5 and slightly longer than article 4; all articles except dactylus with sparsely spaced plumulose spines. Pereopod 6 (Fig. 3): Similar to pereopod 5 but about 10% longer; coxal gill smaller only slightly longer than % article 2; coxa with small anterior lobe and no posterior lobe but with small posterodistal projection; article 6 relatively longer than for pereopod 5. Pereopod 7 (Fig. 3): 10% longer than pereopod 6 and longest pereopod; coxal gill absent; coxa semilunate about twice as wide as long; article 2 proximally expanded and distally narrowed; articles 4 and 5 subequal in length about % length article 6; other- wise similar to pereopod 6. Pereon: Without dorsal or lateral projections. Pleon (Figs 1 and 4): Pleonal epimera with lateral ridge, 1-2 with small (3 with large) posteroventral tooth, dorsally smooth, all with minor serrations on ventral margin and PROG. LINN. SOC. N.S.W., 110 (4), (1988) 1989 BaD A NEW SPECIES OF MELITA Fig. 3. Melita plumulosa sp. nov. Holotype male; scale bar = 0.2mm. See legend to Fig. 1 for key to abbreviations. PROG. LINN. SOC. N.S.W., 110 (4), (1988) 1989 W. ZEIDLER BOS 2-3 with small spine near anteroventral corner; pleonite 4 bearing large dorsal tooth projecting posteriorly to limit of pleonite 5; pleonite 5 with one dorsolateral tooth on each side accompanied by one dorsally placed, upwardly curved spine; pleonite 6 gently rounded posteriorly with dorsal depression near base of telson and acute lower posterior corner. Uropod 1 (Fig. 1): Reaching to % length of outer ramus of uropod 3; rami subequal slightly shorter than peduncle each tipped with three large and two small spines; inner and outer margins of peduncle and rami spinulated; peduncle with large spine at in- sertion of outer ramus and proximally on lower margin almost % along length of peduncle. Uropod 2 (Fig. 1): Shorter than uropod 1; outer ramus slightly shorter than inner, both armed similarly to uropod 1; peduncle as long as inner ramus, inner and outer margins with three large spines. Uropod 3 (Fig. 1): Greatly elongate; outer ramus about twice length peduncle, bearing bunches of spines along both margins, with spine-like distal article surrounded by five smaller spines; inner ramus short, scale-like about 4% length peduncle with one large subterminal spine; peduncle with one small and one large spine mid-ventrally, four spines on posteroventral corner, two spines on outer posterodorsal corner and three spines near inner posterodistal corner. Telson (Fig. 1): About % or less length peduncle uropod 3, cleft to base, outer margins rounded, inner margins slightly excavate for terminal %; each lobe with two large sub- apical spines near inner and outer margin on dorsal surface. DESCRIPTION OF ALLOTYPE: Ovigerous female 4.7mm (AM, P37290) with eleven eggs in brood pouch. Differs from male as follows: Setae and spines of appendages not plumulose. Antenna 1 (Fig. 5): Setae not as long or as numerous; accessory flagellum 2-articulate plus tiny third article; flagellum with 17 articles. Antenna 2 (Fig. 5): Gland cone relatively larger, reaching % along peduncular article 3; peduncular article 4 only marginally longer than article 5; flagellum with 7 articles together slightly longer than peduncular article 4. Coxae: Slightly more elongate. Gnathopod 1 (Fig. 5): Article 6 without humps; dactylus as long as palm. Gnathopod 2 (Fig. 5): Only slightly larger than gnathopod 1; article 6 slightly longer than article 4 and 5 combined, palm oblique undefined by teeth or processes; dactylus not over-riding inner face of palm. Pereopod 6 (Fig. 5): Coxa with relatively long anterior lobe almost as long as rest of coxa. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 334 A NEW SPECIES OF MELITA Fig. 4. Melita plumulosa sp. nov. Holotype male; scale bars = 0.1mm and ().2mm respectively. See legend to Fig. 1 for key to abbreviations. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 W. ZEIDLER 335 Oostegites (Fig. 5): On gnathopod 2 and pereopods 3-5, very slender with sparse margi- nal setae, all similar in shape and size. Uropod 1: Reaching less than % length outer ramus uropod 3. Paratypes: 9 males, 4 females (SAM, C 4076) show little variation from the holotype or allotype. In some specimens pleonal epimera 2 and 3 have two anteroventral spines. The spines and setae of the males are not as plumulose as in the holotype and of the four females one is ovigerous with spines and setae like the allotype, the other three have some spines and setae plumulose as in the males. Etymology: The name refers to the irregularly plumulose nature of the spines and setae. DISCUSSION M. plumulosa is only known from the type locality. “The Blue Pools’ at Angourie were formed as a result of quarrying in the 1920s, the resultant excavation having filled with freshwater from groundwater discharge and surface run-off. Thus colonization has taken place recently and one would expect to find the species in coastal rivers nearby. There are two main lakes at Angourie both less than 100m from the sea but amphipods were only found in the northern one. This lake is the smaller of the two, about 20m in diameter and about 8m deep. Although the habitat is very close to the sea and would be affected by salt spray the surface water is always fresh (C. Creighton pers. comm.) but stratification may occur with brackish water being kept near the bottom. A distinctive feature of MM. plumulosa is the irregularly plumulose character of the spines and setae, more developed in males, and common to all parts of the animal including the mouthparts. I have been unable to find this character in any of the species of Melita that I have examined nor could I find a reference to it for any other species of Melita or indeed for any species of gammaridean amphipod. Another unusual feature is the humps on article 6 of the first male gnathopods, a character which appears to be rare amongst melitids and apart from M. awa Barnard, 1972, does not occur in any other species although some such as M. hergensis Reid, 1939, and M. palmata (Montague, 1804) have the first male gnathopods with the anterodistal margin broadly rounded to overlap the base of the dactylus. M. plumulosa is most similar to the New Zealand species M. awa, particularly in the form of the gnathopods. However, apart from several minor differences, M. awa is dis- tinguished from M. plumulosa by the lack of a dorsal spine on pleonite 4; in that the posteroventral angle of epimeron 3 consists of a small tooth and that the telson is slightly longer than half of the peduncle of uropod 3. In Australia this species must be compared with M. festzva (Chilton, 1885) with which it shares the same pleonal tooth formula. M. festiva differs from M. plumulosa in several characters but mainly in (1) the shape of article 6 of the male gnathopod 1 which lacks humps and is about twice as long as wide; (2) the large size of article 6 of the male gnathopod 2 which is longer than the rest of the limb and also the distinct teeth on the palm; (3) the relatively short, blunt dactylus of the male gnathopod 2 which arises from article 6 off-centre; (4) the irregularly toothed palm of the female gnathopod 2; (5) the small posteroventral tooth of epimeron 3; and (6) the telson length which is equivalent to the peduncle of uropod 3. Other species (none Australian) which also share the same pleonal tooth formula are M. abyssorum Stephenson, 1944, M. hergensis, M. palmata and M. planaterga Kunkel, PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 336 A NEW SPECIES OF MELITA Fig. 5. Melita plumulosa sp. nov. Allotype female; A and B, oostegite from G2 and P5 respectively; scale bar = Q).2mm. Sec legend to Fig. 1 for key to abbreviations. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 W. ZEIDLER 338i 1910. These species differ from M/Z. plumulosa in the following main characters. In M. abyssorum the eyes are absent and the male gnathopod 2 has a large palmar tooth. In /. hergensis article 6 of the male gnathopod 1 has the anterodistal margin broadly rounded to overlap the base of the dactylus and the inner surface of the palm of gnathopod 2 is finely setose. In MM. palmata the male gnathopod 1 is similar to M. hergensis as is the setation of gnathopod 2 but in addition article 6 is very broad distally. In M. planaterga the male gnathopod 1 is similar to M. hergensis, the male gnathopod 2 is very setose, arti- cle 2 of pereopods 5-7 is inflated and rounded and the posteroventral angle of epimeron 3 consists of a medium tooth. It is worth noting here that Barnard (1962) lists MZ. planaterga as a species without dorsal epimeral spines however Karaman (1981) clearly illustrates epimeral spines similar to those of M. plumulosa as do Lazo-Wasem and Gable (1987) who redescribed M. planaterga from recently-discovered type material, although they regard Karaman’s specimen as a different and possibly undescribed species. Apart from the pleonal tooth formula and the absence of humps on article 6 of the male gnathopod 1, other Australian species differ from M. plumulosa principally in the following manner. In M. matilda Barnard, 1972 (1) the accessory flagellum of antenna 1 consists of 5 articles; (2) the palm of the male gnathopod 2 has a deep hollow with spines to accommodate the dactylus; (3) the female coxa 6 is unmodified; (4) the posterodistal angle of article 2 of pereopods 5-7 is not produced; (5) the setae and spines on uropod 3, and at least some on articles 4-6 of pereopods 5-7, are about twice as long; (6) mandi- bular palp article 3 is longer than article 2. In M. oba Barnard, 1972 (1) the accessory flagellum of antenna 1 consists of 4-5 articles; (2) the medial face of the male gnathopod 2 is very setose and there is no hollow for the dactylus which only partly over-rides the palm; (3) article 4 of the female gnathopod 2 is without a sharp process; (4) coxa 4 1s expanded ventrally and slightly excavate; (5) article 2 of pereopods 5-7 is more regularly ovate with rounded postero-distal lobes; (6) the posteroventral angle of epimeron 3 is only slightly produced; (7) mandibular palp article 3 is longer than article 2. In M. zeylanica Stebbing, 1904 (1) the medial face of the palm of the male gnathopod 2 is very setose; (2) coxa 4 is deeply excavate posterodistally; (3) article 2 of pereopods 5-7 is inflated and rounded; (4) the posteroventral angle of epimeron 3 consists of a small tooth. NOTE ON THE GENUS ABLUDOMELITA KARAMAN, 1981 In his study of the genus Melita, Karaman (1981) determined two groups of species, those ‘without dorsal oblique row of setae on inner lobe of maxilla 2 and prevalently with 1- segmented outer ramus of uropod 3’ and those ‘with dorsal oblique row of setae on inner lobe of maxilla 2 and prevalently with 2- segmented outer ramus of uropod 3° Karaman considered these characters to be of generic significance and consequently erected the genus Abludomelita to accommodate the latter group, the former being retained in the genus Melita. In M. plumulosa the inner lobe of maxilla 2 is without a dorsal oblique row of setae and the outer ramus of uropod 3 can be considered 2-segmented. Thus following Kara- man it could be placed into either of the above two groups. Based on the character of maxilla 2 alone M. plumulosa would fit into the Melita group but this would deny its close relationship to M. awa which is placed in the Abludomelita group by Karaman. As the two characters on which Abludomelita is based are plesiomorphic the genus 1s at best paraphyletic and probably polyphyletic. The setation of maxilla 2 has not been critically examined for most species of Melita and the degree of variation is unknown. Also I do not consider the loss of article 2 on the outer ramus of uropod 3 to be of generic PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 338 A NEW SPECIES OF MELITA significance in Melita as this article can be so reduced in some species that it is hidden by, or is indistinguishable from, adjacent spines (i.e. when is an article just another spine?). Karaman admits that the ‘pilosity of maxilla 2 is not described in some species’ and that some of his conclusions are based solely on the form of uropod 3. Clearly Melita needs much more detailed study before new generic concepts are put forward. In consideration of the above I am reluctant to accept Karaman’s division of Melita and his new genus Adludomelita until a more detailed revision of the genus 1s undertaken. In the meantime I am referring my species to the genus Melita. Of the species mentioned in this paper Karaman transferred the following to Abludomelita: M. abyssorum, M. awa, M. festiva, M. matilda and M. oba. ACKNOWLEDGEMENTS I wish to thank Mr C. Creighton of Landwater Management, Angourie, and Dr B. V. Timms, Avondale College, Cooranbong, N.S.W. for information regarding the type locality. Dr W. F. Ponder, The Australian Museum, Sydney, identified the hydrobiid snail. Dr J. K. Lowry, The Australian Museum, Sydney, kindly reviewed the manuscript and provided comparative material for this study. References BARNARD, J. L., 1962. — Benthic marine Amphipoda of southern California: Families Tironidae to Gammaridae. Pacific Nat. 3: 73-115. ——., 1972. — Gammaridean Amphipoda of Australia, Part I. Smithson. Contrib. Zool. 103: 1-333. , and BARNARD, C. M., 1983. — Freshwater Amphipoda of the World. 2 Vols. 830 pp. Mt Vernon, Virginia: Hayfield Associates. COoL_LetTT, L. C., COLLINS, A. J., GipBs, P. J., and WEsT, R. J., 1981. — Shallow dredging as a strategy for the control of sublittoral macrophytes: a case study in Tuggerah Lakes, New South Wales. Aust. J. Mar. Freshw. Res. 32: 563-571. Kanaas, M. I., and GEDDES, M. C., 1984. — The effects of salinity on the distribution of amphipods in The Coorong, South Australia, in relation to their salinity tolerance. Trans Roy. Soc. S. Aust. 108(3): 139-145. KARAMAN, G. S., 1981. — Redescription of Melita planaterga Kunkel 1910 from Bermuda islands with revision of genera Melita Leach and Abludomelita n. gen. Poljoprivreda i Sumarstvo, Titograd 27(1): 29-50. LAzO-WASEM, E. A., and GABLE, M. F., 1987 — A review of recently discovered type specimens of Bermuda Amphipoda (Crustacea: Peracarida) described by B. W. Kunkel (1882-1969). Proc. Biol. Soc. Wash. 100(2): 321-336. PooRE, G. C. B., 1982. — Benthic communities of the Gippsland Lakes, Victoria. Aust. J. Mar. Freshw. Res. 33: 901-915. PoTTER, I. C., LENANTON, R. C. J., LONERGAN, N., CHRYSTAL, P., CAPUTI, N., and GRANT, C., 1981. — The fish and crab fauna of the Peel- Harvey estuarine system in relation to the presence of Cladophora. Bull. No. 88. Dept. Cons. Env. W.A. SHEARD, K., 1937. — A catalogue of Australian Gammaridea. Trans Roy. Soc. S. Aust. 61: 17-29. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 Baseline Survey of the Benthic Macrofauna of Twofold Bay, N.S.W., with a Discussion of the Marine Species introduced into the Bay PAT HUTCHINGS, JOS VAN DER VELDE, and STEVE KEABLE HUTCHINGS, P. A., VAN DER VELDE, J. T., and KEABLE, S. J. Baseline survey of the benthic macrofauna of Twofold Bay, N.S.W., with a discussion of the marine species introduced into the Bay. Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 339-367. A baseline survey of the benthic macrofauna of Twofold Bay, southern N.S.W., was carried out during 1984-85. Sampling was concentrated in intertidal and shallow sub- tidal habitats. Samples were collected seasonally and, although sampling was largely qualitative, some attempts were made to assess the abundance of species and this is indicated in the species list given. The survey originated in response to a study on ballast water which is regularly discharged into Twofold Bay (Williams et a/., 1988). The ballast water contained living organisms and Williams ef al. (1988) suggested that these animals could survive discharge into the Bay. The purpose of this survey was to conduct a baseline study of the macrofauna of Twofold Bay and to determine if any non-indigenous species had been introduced into the bay. Seven introduced species were found and the possible methods by which these became established in the bay are discussed. P. A. Hutchings, J. van der Velde and S. J. Keable, The Invertebrate Division, Australian Museum, Sydney, Australia 2000; manuscript received 30 June 1987, accepted for publication 20 July 1988. INTRODUCTION Background to Study During the 1970s considerable concern was expressed both overseas and in Aus- tralia on the likelihood of marine introductions via ballast water discharged into ports. Subsequent colonization and the potential for the development of them to become ‘pest’ species was also of concern (Friese, 1973; Medcof, 1975; Medcof and Wolf, 1975). Carlton (1985, 1987) has recently reviewed species introduced via ballast water on a world wide basis. In response to this concern primarily within the fishing industry, the then New South Wales Department of Fisheries, now the Department of Agriculture and Fisher- ies, undertook a survey of the ballast water of Japanese ships discharging ballast water into selected Australian ports. Their survey revealed a variety of living organisms in the ballast water belonging to a number of groups, including 8 non-indigenous species (6 copepods, 1 mysid and 1 amphipod) and an additional 14 species of copepods and 4 non copepod taxa with an Indo-Pacific distribution. Twenty one copepods and 22 other species, from the ballast water sampled, could not be identified to species. Sediment in the bottom of the ballast water tanks contained 8 non-indigenous species, 8 cosmopoli- tan and 27 other taxa not identified (Williams et al., 1988). They determined by experi- ments that such organisms could survive discharge through the pumps into the water and thus could possibly settle and establish populations in an Australian port. Twofold Bay on the south coast of New South Wales (37°05’S, 149°54’E) is one of the ports identified by Williams e¢ al. (1988) as receiving ballast water containing living organisms on a regular basis from northern Japan during loading of woodchips at the wharf on the southern shores of the bay. Medcof (1975), in an earlier survey had also found living ostracods, crustacean larvae, adult and larval polychaetes and chaeto- gnaths in ballast water of a woodchip carrier at Eden, Twofold Bay. PROC. LINN. SOC. N.SW,, 110 (4), (1988) 1989 340 BENTHIC MACROFAUNA OF TWOFOLD BAY Twofold Bay has been a port since the late 1800s, for a variety of goods including whale products, gold, agricultural produce and most recently for woodchip (Matthews, 1947). Thus considerable scope has existed for introductions via fouling organisms on the bottoms of ships and more recently via ballast water (Medcof 1975; Williams et al., 1988). The marine fauna of the bay has not however been documented. During 1984 and 1985, The Australian Museum undertook a baseline study of Twofold Bay as a follow-up to the survey of Williams et a/., 1988. The aims of the survey were (1) to document the macrobenthic fauna of the area, and (2) identify any marine in- troductions. Any species recognized as non-indigenous to the area, could then be the subject of a subsequent study, to assess what impact, if any, this species is having on the natural fauna and recommend control measures. This baseline study will also facilitate the documentation of any subsequent introductions. Curalo Lagoon 3 CALLE CALLE BAY ae: 2) EDEN ey Quarantine Bay Shadracks Creek Murrumbulga Pt. TWOFOLD BAY NULLICA BAY Sm Nullica River EAST BOYD BAY unganno rm LP ; Fisheries Creek Fig. 1. Map showing location of sampling sites within Iwofold Bay. Study Area Twofold Bay is a large open bay, consisting of 3 smaller bays, Calle Calle, Nullica and East Boyd Bay (Fig. 1). The headlands and corresponding rock platforms are of PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VAN DER VELDE ANDS. J. KEABLE 341 Devonian sedimentary strata. The beaches between these bays are mini-barrier dune systems which form Curalo Lagoon and are also evident at the mouth of Fisheries Creek, Towamba and Nullica Rivers. Rationale for sampling strategy Ballast water is usually taken on board after unloading the cargo. Ports are usually sited in estuarine or shallow protected bays so that the fauna taken on board with the ballast water is primarily permanent estuarine plankton and planktonic larval stages of estuarine or shallow water species. The adults of such species tend to live as encrusting organisms or benthic organisms in soft sediment or seagrass communites, therefore sampling was concentrated in these habitats. TABLE 1 A Breakdown of the various Types of habitats sampled at each Site sheltered yearns Murrumbulga Pt marine ene GEGONTEl saa ca caads cao cc Munganno Pt intertidal estuanine(OySters)) tae ere eee ere eer Nullica River Fisheries Creek natural va Pct sheltered waver Murrumbulga Pt Hard substrata eS Coe EXPOSEd iy eee ete Munganno Pt aneticial —whariepilesmbreakwate tern rs it Treaties neta raoeera ae Munganno Pt Quarantine Bay shallow Posidonia (seagrass beds) .......... Quarantine Bay marine Zi Ss deep (10-20m) — pipe dredge sampling marine sand Soft substrata (infauna) lasoontwlu pp7al(Scagnass; beds) maa rrt- nc acr acre rseie ramen ie Fisheries Creek estuarine < tidal’ Zosteral(Sea grassibeds) eri areas Serer eer een ai Nullica River Curalo Lagoon Shadracks Creek Sal trivars hie a westdirsceakacerns pm ctennte cs waves Gatte pease ceienerecsy auc rae Fisheries Creek Curalo Lagoon METHODS The habitats sampled in Twofold Bay (Table 1) were: — a. Fouling communities on wharfs and piles. Seagrass communities at the mouth of rivers and creeks. c. Soft bottom communities and intertidal encrusting fauna in shallow estuarine areas. Intertidal rocks around the loading wharf at the woodchip mill. e. Saltmarsh areas. No attempt was made to sample the pelagic or planktonic communities within the Bay. Fig. 1 indicates the location of each of the sampling sites. Details of sampling and the number of samples collected in each habitat are given in Table 2. A variety of qualitative collecting techniques were used to sample the fauna, although in the Appen- dix some indication of abundance 1s given for each species. Details of collection methods appear in Hutchings et al. (1986a, b). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 342 BENTHIC.MACROFAUNA OF TWOFOLD BAY TABLE 2 Details of Times of Sampling at each Site and the Number of Samples collected in each Habitat, during each Collection Period Unit of collection = 36cm x 44cm plastic bag Variation between months in collection unit number caused by bad weather and time available for collection SAMPLE REPLICATES COLLECTION UNITS SITES YEAR 1984 1985 MONTH SEPT DEC MAR JUN SEPT DEC Munganno Point Intertidal rock platform 3 2 2 3 3 4 Subtidal rock platform 3 1 1 2 1 2 Wharf piles 3 2 3 3) 3 5 Airlift sediment 1 1 1 — _ — (a) Fisheries Creek Intertidal rocks = 1 1 1 1 1 Sand sievings (intertidal)* 6 6 6 6 6 6 Mud sievings (intertidal)* 6 6 6 6 6 6 Mud sievings, Ruppia* 6 6 6 6 6 6 Saltmarsh 6 1 1 1 1 1 Nullica River Sand sievings (intertidal)* = 6 6 6 6 6 Mud sievings (intertidal)* 6 6 6 6 6 6 Mud sievings, Zostera* 6 6 6 6 6 6 Intertidal rocks 2 1 2 1 1 2 Shadracks Creek Mud sievings, Zostera* 6 6 6 6 6 6 Sand sievings (intertidal)* 6 — — — — — (a) Murrumbulga Point Intertidal rock platform 2 2 3 3 3 3 Subtidal rock platform 5 2 3 3 3 2 Quarantine Bay Airlift sediment (Poszdonia) 4 4 4 4 4 4 Amphipod trap, wharf 1 — 1 1 1 1 Curalo Lagoon Sand sieving, Zostera* 6 6 6 6 6 6 Saltmarsh 1 1 1 1 1 1 Total 79 66 71 71 70 74 * Hand corer used. (a) Discontinued. On one occasion, permission was given by the shipping authorities to collect the accumulated sediment in the bottom of the ballast water tanks. Salinity was measured during each collecting period for the estuarine areas sampled. a. Description of Sampling Sites and Salinity Regimes 1. Curalo Lagoon A shallow brackish water lagoon behind the most northern beach in Twofold Bay. It was Closed to the sea in July 1984 for a short period and during this time the lagoon was virtually fresh and the bar was breached between July and September 1984. The lagoon PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VAN DER VELDE AND S. J. KEABLE 343 then remained open to the sea for the rest of the survey and the salinity ranged for 20- 32°/oo, with an average of 26°/o0. Zostera seagrass beds were present near the entrance; they replaced Ruppia seagrass beds present before the salinity increased. An extensive saltmarsh occurs on the eastern margin. 2. Murrumbulga Point A south-east facing sheltered rocky shore with the intertidal rock platform approxi- mately 15-20m wide, which has been eroded flat, with scattered rubble, boulders and sand occurring at the north-eastern end. Zonation is not as apparent as at Munganno Point, described below, because of its more sheltered nature, gentle slope and the presence of numerous tidal pools. The subtidal rock ‘platform’ is boulder-strewn and uneven, extending gradually to a depth of 9m. The boulders have a cover of kelp and other algae, while the crevices are dominated by sea urchins and encrusting red calcareous algae. 3. Quarantine Bay This bay is protected from heavy seas by a breakwater. Patchy Posidonia seagrass beds begin at approximately 2m depth between the rocks and sand. Encrusting and cryptic fauna were also collected from these rocks. The pylons of the recreational/public boating wharf and the intertidal breakwater wall were examined. 4. Shadracks Creek A brackish tidal creek with sandy banks and Zostera seagrass growing on a rubble covered mud bottom. It forms a small lagoon behind Legges Beach. It had an average salinity of 14.5°/oo (range 0-27°/oo) and was closed in March, 1985, and remained closed for the rest of the sampling period. 5. Nullica River The estuary has extensive intertidal mud and sand flats. Zostera seagrass grows in the channel and tide pools and oysters cover the intertidal rocks. It was always open to the sea at times of sampling with an average salinity of 21.7°/00 (range 11-30°/o0). 6. Fisheries Creek A brackish tidal creek with intertidal mud and sand flats, oyster covered rocks, and the seagrasses Zostera and Ruppia growing in the deeper back waters. Saltmarshes are present along the bank. The creek showed an average salinity of 29.8°/oo (range of 20-36°/o0), with salinity falling steeply from the mouth to the seagrass beds, a distance of approximately 500m. The creek was closed to the sea in March 1985 and in full flood in December later that year. 7. Munganno Point A north-west facing exposed rocky shore with the intertidal rock platform approx1- mately 4m wide, backed by a small cliff. The platform varies from a steeply sloping rock face to loose piled rubble with a few tide pools. There is some evidence of zonation. The lower zone is dominated by cunjevoi (Pyura stolonifera (Heller) ) and mat-forming weed. The upper zone is dominated by serpulid worms (Galeolaria caespitosa Savigny) and barnacles. The subtidal rock platform is a tiered ‘patchy’ outcrop of rocks which extends down to sand at approximately 6m depth. The rocks support a healthy cover of kelp (Ecklonia radiata (C.Ag.) J.Ag). PROC. LINN. SOC. N.S.W,, 110 (4), (1988) 1989 344 BENTHIC MACROFAUNA OF TWOFOLD BAY The wharf at Munganno Point extends from the shore for 244m, with maximum water depth 15m. The pylons are constructed of concrete-encased steel. Pylons close to the shore are thickly encrusted with calcareous serpulid worm tubes intertidally, whilst supporting a heavy growth of tunicates, sponges and kelp for their entire length below the low water mark. RESULTS In the Appendix the macrobenthic fauna collected at each locality is given, together with an indication of abundance and frequency. Many of the polychaetes and small crustaceans have been identified only to the generic level, as they represent either new species or new records for the area. Representatives of each taxon have been lodged at the Australian Museum and assigned registration numbers to facilitate further identifi- cation or confirmation. The following species have been introduced and are non-indigenous to the bay: Styela plicata (Ascidian); Polycera capensis, Theba pisana and Crassostrea gigas (Molluscs); Notomegabalanus algicola, Carcinus maenas and Eurylana arcuata (Crustaceans). Each of these is discussed in detail below. Styela plicata (Lesueur) This ascidean or sea squirt has long been known to occur on Australian shores. We recorded it from the subtidal rocks amongst the Poszdonia seagrass in Quarantine Bay. It has a wide distribution within Australia and is found in many ports of the world. Recent analysis of the biogeography of this species has indicated that it is non- indigenous to Australia (Kott, 1985). Styela plicata is a recognized fouling organism and is likely to have been introduced to Australia attached to ships’ hulls. The earliest Australian records date from 1878. The origin of the Twofold Bay colony is unknown. It may have resulted from the spread of populations already established in Australia or as a direct import from over- seas shipping. This could have occurred at any time during the 150 years that the port of Twofold Bay has existed (Matthews, 1947). Theba pisana (Muller) This salt-tolerant terrestrial snail was collected from the terrestrial margins of the saltmarsh at Curalo Lagoon. It is recognized as a pest of agricultural crops and gardens (Smith and Kershaw, 1979). Baker (1986) has reviewed the introduction and spread of 7? pesana in Australia. It first appeared in South Australia prior to 1928. It is now widely distributed over most of southern Australia but appears to favour a mediterranean-type climate. Its previous dis- tribution includes Europe, the Mediterranean, North Africa, Atlantic Isles and British Isles. It has also been introduced to North America and South Africa. The mode of introduction into Australia is not known, but it may have been brought to Australia as a food item by Italian migrants (P. Colman, pers. comm.). Crassostrea gigas (Thunberg) The Pacific oyster was recorded, in small numbers, at Twofold Bay on the intertidal rocks at the mouth of the Nullica River. This is the first record of occurrence in Twofold Bay (IT. Mundy, pers. comm.). Shipments of the Pacific oyster were made from Japan in 1947-8 to establish popu- lations in southern Tasmania (Thomson, 1952). In 1955, adult Pacific oysters from Tas- mania were transported to Mallacoota, Victoria, just south of Twofold Bay. By 1958 a PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VANDER VELDE ANDS. J. KEABLE 345 quarter of the population was still alive, but no spatfall was observed during this time (Thomson, 1959). Wolf and Medcof (1973-4) documented the distribution of Crassostrea gigas in New South Wales. They provide an accurate documentation of all the Australian introduc- tions and subsequent dispersal of stock. Although they did not record the species from Twofold Bay, the oyster was recorded from Pambula (1967) and Merimbula (1973) just to the north. Subsequent papers (Medcof and Wolf 1975; Holliday and Nell, 1985; Cole- man, 1986) have further examined the expansion of the range of Crassostrea gigas and dis- cuss the problems this oyster has caused in the N.S.W. oyster industry which has traditionally been based on the Sydney rock oyster Saccostrea commercialis. A breeding population of C. gigas has established itself in Port Stephens, N.S.W. (Holliday and Nell, 1985). The Pacific oyster is providing a case study of the effects of a ‘pest’ on an established fishery. Polycera capensis (Quoy and Gaimard) The first Australian records of this opisthobranch mollusc date from 1927, the animals being collected from Sydney Harbour and described by Allan (1932) as a new species Polycera conspicua. Thompson (1975) reported that the species described by Allan (1932) had been synonymized with Polycera capensis (Quoy and Gaimard) a species commonly found in, and first described from, South Africa. Thompson (1975) also noted that further speci- mens had been taken from Sydney Harbour and Botany Bay, N.S.W., where the species was at least seasonally abundant. Burn (1978) gives the Australian distribution of P capensis as the area between Broken Bay and Kiama, N.S.W. Willan and Coleman (1984) repeated the Australian distribution given by Burn (1978) and state ‘P. capensis was probably introduced to Australia by shipping’. A single specimen was collected in December 1985 by us from Twofold Bay, at the subtidal rocks Munganno Point, adjacent to the woodchip loading wharf. Specimens were also taken from this area by Rudman (pers. comm.) in March, 1986. The evidence to date suggests that P capensis has been introduced into Australia from South Africa, by ship fouling. It may have been introduced to Twofold Bay either directly in this manner or from the dispersal of populations established in other Australian ports. Polycera capensis was not recorded by us as an introduction to Twofold Bay in our original report (Hutchings et al., 1986b) as the literature relating to it has only recently been brought to our attention. Notomegabalanus algicola (Pilsbry) This barnacle was originally described from South Africa and was first noted in Australia by Pope in 1943 (see Allen, 1953). It seems likely that the first introduction occurred in the Sydney region, although the barnacle may have been introduced into New South Wales on several occasions. Allen (1953) records this barnacle from Eden to Port Stephens and suggests that it was transported to Australia as a fouling species on the bottom of ships. Allen states that during the late 1940s to early 1950s it rapidly increased in numbers and became one of the most common sublittoral barnacles on the open coast. The record of this barnacle from Twofold Bay is therefore not surprising. It was found to be very common on the intertidal rocks and wharf piles of Munganno Point. PROG. LINN. SOG. N.S.W., 110 (4), (1988) 1989 346 BENTHIC MACROFAUNA OF TWOFOLD BAY Carcinus maenas (Linnaeus) This species, also known as the European swimming crab, was first recorded from Australia by Fulton and Grant (1900; 1901) from Port Phillip Bay, Victoria. They suggest it was introduced amongst the dense fouling growth on the bottoms of wooden ships and that the young crabs could easily live amongst this growth during the long sea voyage from Europe. This crab has since been recorded from Westernport Bay to Mallacoota (Allen, 1953), and Healy and Yaldwyn (1970) recorded it as quite abundant on parts of the Vic- torian coastline. Day and Hutchings (1984) tentatively recorded it from Merimbula, New South Wales, and specimens with collection localities in the vicinity of Narooma, New South Wales, are held at the Australian Museum. We have found it commonly around [wofold Bay, amongst intertidal rocks as well as intertidal mud. Our survey thus confirms the continued extension of C. maenas populations to the north. Recent intro- ductions of C. maenas have also been reported in South Australia, and a single specimen has been found in Western Australia (Zeidler, 1978; Rosenzweig, 1984). To date, no information is available on the impact of this introduced species on our native populations. Overseas experience indicates that such an ‘aggressive predator constitutes a potential threat to native marine fauna (Joska and Branch, 1986). Eurylana arcuata (Hale) This cirolanid isopod is well known only from New Zealand and Chile, South America. It is believed to have been a recently-introduced species in San Francisco Bay, U.S.A. (Bowman eé al., 1981). Bowman et al. suggest the isopod could have been intro- duced to San Francisco Bay in this case either in fouling, such as that found on propeller-shaft housing, or via ballast water. They gave three localities in Australia where this isopod has been previously reported (Port Jackson and Broughton Island, New South Wales and Port Willunga, South Australia) and suggested that it may have been introduced at these points. Bruce (1986) has recorded a single specimen from New- castle harbour and Day and Hutchings (1984) recorded the species tentatively from Pambula and Merimbula. At Twofold Bay, the species occurred on the intertidal rock platform at Murrumbulga and offshore in deep water sediments. This disjunct distribution within Australia, associated with port facilities, and the fact that Eurylana arcuata is not recognized as an active swimmer (N. Bruce, pers. comm.), indicates that it may have been introduced on several different occasions at different localities. Sediment in ballast water tanks The sediment collected from the bottom of the empty ballast water tanks contained one juvenile crab and 60 copepods. The crab was identified as Charybdis cf. feratus which has an Indo-Pacific distribution from Japan and East Asia to the east coast of Africa and south to Victoria, Australia. No specimens of this species were recorded during the sur- vey and we cannot be certain that the crab was alive when the mud was collected although it was well preserved and its presence in the sediment confirms the findings of Williams et al. (1988). The 60 copepods present in the mud have not been identified further. However the material has been deposited in the Australian Museum. DISCUSSION The benthic macrofauna of Twofold Bay is a rich and diverse one, with over 570 taxa collected. This number includes 13 species of demersal fish which normally inhabit PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VANDER VELDE ANDS,. J. KEABLE 347 the seagrass beds which were sampled for their benthic invertebrates. This level of rich- ness appears high for southern New South Wales and exceeds the 246 invertebrate species identified at nearby Merimbula (Day and Hutchings, 1984). There is some over- lap in species composition in the comparable habitats sampled at these two localities. Of the seven marine species recorded during this survey as non-indigenous to Twofold Bay, two may have been introduced via ballast water, Eurylana arcuata and Crass- ostrea gigas. The remaining species, with the exception of Theba pisana, were probably in- troduced as fouling organisms. The influence of these seven introduced species on the ecology of the indigenous marine life in the bay is not known. ‘To date only the Pacific oyster Crassostrea gigas has been recognized as posing a possible commercial threat to fish- eries and this is a state wide problem not one restricted to Twofold Bay. The spread of Carcinus maenas along the southern and south-eastern Australian coastline and possibly into Western Australia may warrant further studies including ones to assess the impact this species has on the local fauna. Joska and Branch (1986) describe Carcinus maenas as an ‘aggressive predator’ where ‘the great strength of its nippers and its aggressive, non-selective predatory habits have made it a pest in at least one of its adopted habitats. They report C. maenas feeding on the clam-beds on the New England coast of the U.S.A. This survey provides a basic inventory of the macrofauna of shallow intertidal and subtidal benthic communities in Twofold Bay and should allow any subsequent intro- ductions to be identified and the timing of the introduction to be assessed. To date we believe that the evidence for introductions via ballast water into Twofold Bay are only tentative and the effect on the native fauna not well documented except in the case of Crassostrea gigas. Suggestions made by Paxton and Hoese (1985) and Williams et al. (1988), that all ballast water being discharged into Australian ports be treated in order to kill any living organisms before the water is discharged, are at this stage we believe premature. The problems of introducing the necessary legislation which would involve co-operation between several countries, and the enforcing of any such legislation are such that far more documentation of introductions via ballast water and their impact on the native fauna is needed for their justification. Carlton (1985, 1987) in a recent review of ballast water introductions summarizes those attributed to ballast water but in many cases the data substantiating such a mechanism of introduction is fairly tenuous. Instead of pursuing costly legislation to sterilize ballast water, baseline surveys of the fauna and flora of those Australian ports which regularly receive ballast water (especially those which receive ballast water from ports subjected to similar water tem- perature regimes as those in the Australian port) should be conducted. Such a monitoring programme and the identification of any introduction and documentation of the impact these introductions are having on the natural fauna will provide the necessary data which would be necessary before any legislative programme could even be considered and drawn up and enforced. ACKNOWLEDGEMENTS The study was funded by a Fishing Industry Research Trust Account grant (84-49). The work would not have been possible without the assistance of the following people: E. A. Bamber, L. M. Walker, A. C. Paul, A. L. Reid, S. J. Perry and P. L. Albertson of the Australian Museum who helped in the field; P. Albertson, L. Albertson, L. Matthews and D. Winn of the Australian Museum who helped in the laboratory; P. Went (General Manager) and G. Hodkinson (Shipping Manager) of Harris-Daishowa (Aust.) Pty. Ltd. for access to sample the Munganno Point area and for helpful co-operation shown at all times; D. Laffam (Customs Office at Eden) for his help in organizing the collection of PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 348 BENTHIC MACROFAUNA OF TWOFOLD BAY ballast-tank mud from a Malayasian carrier in September 1984. We are also grateful to the following people for their taxonomic help: Dr H. Paxton, Dr H. ten Hove, Dr K. Fauchauld, Dr P. Knight-Jones, C. Glasby, C. Watson-Russell, L. M. Walker and S. J. Perry (Polychaetes); Dr J. Lowry, Dr N. Bruce, Dr G. Poore, Dr A. Myers, D. Jones, H. Stoddart, P. Berents, H. Tranter, R. Springthorpe, A. Murray, M. Drummond and Dr C. Pregenzer (Crustacea); Dr W. Ponder, Dr W. Rudman, I. Loch, P. Colman, J. Kers- lake, B. Jenkins and T. Cochran (Molluscs); Dr W. Rudman kindly provided us with his unpublished list of Opisthobranchs from Twofold Bay, Dr F. Rowe (Echinoderms and Urochordates); Dr D. Hoese, D. Rennis and S. Reader (fish). Our special thanks go to Dr A. Jones of the Australian Museum for his advice on sampling strategies, Dr N. Tate for the loan of hand corers, A. Reid for the artwork and to P. Dempsey for typing the manuscript. TI. Mundy (Eden’s Fishing Inspector) and R. J. Williams are specially acknowledged for their continued interest, constructive criticism and general assistance throughout the course of the study. References ALLEN, J. K., 1932. — Australian Nudibranchs. Aust. Zool. 7(2): 87-105. ALLEN, F. E., 1953. — Distribution of marine invertebrates by ships. Aust. J. mar. Freshwater Res. 4(2): 307-316. BAKER, G. H., 1986. — The Biology and Control of White Snails (Mollusca: Helicidae), Introduced Pests in Australia. C.S.I.R.O. Australia. Division of Entomology Technical Paper No. 25. 31pp. BowMaN, T. E., BRUCE, N. L. and STANDING, J. D., 1981. — Recent introduction of the cirolanid isopod crustacean Cirolana arcuata into San Francisco Bay. J. Crust. Biol. 1(4): 545-557. BRuCE, N. L., 1986. — Cirolanidae (Crustacea: Isopoda) of Australia. Rec. Aust. Mus. Suppl. 6: 1-239. BuRN, R., 1978. — Anew record of Thecacera pennigera (Montagu, 1815). Opisthobranchia: Polyceridae) from New South Wales. J. Malac. Soc. Aust. 4(1-2): 22. CARLTON, J. T., 1985. — Transoceanic and interoceanic dispersal of coastal marine organisms: the biology of ballast water. Oceanogr. Mar. Biol. Ann. Rev. 23: 313-371. ——., 1987. — Patterns of transoceanic marine biological invasion in the Pacific Ocean. Bull. Mar. Sci. 41(2): 452-465. COLEMAN, N., 1986. — A review of introductions of the Pacific oyster (Crassostrea gigas) around the world and a discussion of the possible ecological consequences of introducing the species into Victoria, Australia. Marine Science Laboratories, Technical Report No. 56. 39pp. Day, J. H., and HUTCHINGS, P. A., 1984. — Descriptive notes on the fauna and flora of Merimbula, Pambula and Back Lakes, New South Wales. Aust. Zool. 21(3): 269-289. FRIESE, U., 1973. — Another Japanese Goby in Australian waters; what next? Koolewong. Proc. Roy. Zool. Soc. N.S.W, 2: 5-7. FULTON, S. W., and GRANT, F. E., 1900. — Note on the occurrence of the European crab, Carcinus maenas, Leach, in Port Phillip. Victorian Nat. 17(8): 147-148. ——.,, 1901. — Some little known Victorian Decapod Crustacea with Description of a New Species. Proc. Roy. Soc. Vict. 14(2): 55-64. HEALY, A., and YALDWYN, J., 1970. — Australian Crustaceans in Colour. Sydney: Reed. HO.u.ipay, J. E., and NELL, J. A., 1985. — Concern over Pacific oyster in Port Stephens. Aust. Fish. 44(11): 29-31. HUTCHINGS, P. A., van der VELDE, J. T., and KEABLE, S. J., 1986a. — Colonisation of New South Wales by non-indigenous marine species. Baseline studies at Twofold Bay, New South Wales. Unpublished final report to the Fisheries Industry Research Committee. FIRTA 84-89. 71pp. , 1986b. — Colonisation of N.S.W. by foreign marine species. Aust. Fish. 45(4): 40-42. JOosKA, M. A., and BRANCH, G., 1986. — The European shore-crab. Another alien invader? Afr. Wildl. 40(2): 63-65. Kort, P., 1985. — The Australian Ascidiacea Part 1. Phlebobranchia and Stolidobranchia. Mem. Qld Mus. 23: 1-440. MATTHEWS, L. J., 1947. — Eden, Twofold Bay. Its Rise, Decline and Return to Prosperity. Port of Sydney Jour- nal 1(3): 89-92. ” MepbcorF, J. C., 1975. — Living marine animals in a ship’s ballast water. Proc. natl Shellfish. Ass. 65: 54-55. , and WoLF, P. H., 1975. — Spread of Pacific oyster worries New South Wales culturists. Aust. Fish. 34(7): 32-38. PAXTON, J. R., and Hoese, D. F., 1985. — The Japanese Sea Bass, Lateolabrax japonicus (Pisces, Percichthyi- dae), an Apparent Marine Introduction into Eastern Australia. Jap. J. Ichthyol. 31(4): 369-372. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VAN DER VELDE AND S. J. KEABLE 349 ROSENZWEIG, P. A., 1984. — A range extension for the European shore crab Carcinus maenas (Linn, 1758) in South Australia. Sth Aust. Nat. 59(1): 18-19. SM!TH, B. J., and KERSHAW, R. C., 1979. — Field Guide to the Non-Marine Molluscs of South Eastern Australia. Canberra: Australian National University Press. THOMPSON, T. E., 1975. — Dorid nudibranchs from eastern Australia (Gastropoda, Opisthobranchia). /. Zool. Lond. 176: 477-517. THOMSON, J. M., 1952. — The acclimatization and growth of the Pacific oyster (Gryphaea gigas) in Australia. Aust. J. Mar. Freshwater Res. 3(1): 64-73. ——., 1959. — The naturalization of the Pacific oyster in Australia. Aust. J. Mar. Freshwater Res. 10(2): 144-149. WILLAN, R. C., and COLEMAN, N., 1984. — Nudibranchs of Australasia. Sydney: Australian Marine Photo- graphic Index. WILLIAMS, R. J., van der WAL, E. J., and KELLY, J., 1988. — Ballast water as a possible vector for the trans- port of non-indigenous marine species. Estuarine, Coastal, Shelf Science 26: 409-420. WOLF, P. H., and Mepbcor, J. C., 1973-4. — Be on your guard against the Pacific oyster. The Fisherman 4(10) BO 24-29. ZEIDLER, W., 1978. — Note on the occurrence of the European shore crab Carcinus maenas (Linn., 1758) in Australia. Sth Aust. Nat. 53(1): 11-12. APPENDIX INVENTORY OF MARINE FAUNA COLLECTED FROM TWOFOLD BAY (In alphabetical order within the major taxonomic groupings) The frequency refers to the rate of recurrence or number of collections in which it was found: Occasionally(o0) 0 D0) molluscs > 40 peracarid crustacea x > 100 other crustacea x > 40) where x represents the number of specimens. The sztes are Curalo Lagoon — 1, Murrumbulga Point — 2, Quarantine Bay — 3, Shadracks Creek — 4, Nullica River estuary 5, Fisheries Creek — 6, Munganno Point — 7, Reconnaissance trip (12-13/7/84) — 8, Deepwater benthic sampling (20-22/2/85) — 9. PROC. LINN. SOG. N.S.W., 110 (4), (1988) 1989 350 BENTHIC MACROFAUNA OF TWOFOLD BAY CLASSIFICATION & DESIGNATION PHYLUM: COELENTERATA Phlyctenactis tuberculosa (Quoy & Gaimard) PHYLUM: PLATYHELMINTHES CLASS: TURBELLARIA: Acoela spp. PHYLUM: NEMERTINEA Nemertean ?sp.1 Nemertean ?sp.2 Nemertean ?sp.3 Nemertean ?sp.4 Nemertean ?sp.5 Nemertean ?sp.6 PHYLUM: ANNELIDA CLASS: POLYCHAETA ORDER: ORBINIIDA FAMILY: ORBINIIDAE Leitoscoloplos normalis Day Scoloplos (Scoloplos) cylindrifer Ehlers Scoloplos (Scoloplos) novaehollandiae (Kinberg) Scoloplos (Scoloplos) simplex (Hutchings) FAMILY: PARAONIDAE ORDER: SPIONIDA FAMILY: SPIONIDAE Aonides oxycephela (Sars) Australospio trifida Blake & Kudenov Boccardta chilensis Blake & Woodwick Boccardiella sp. Carazziella victoriensis Blake & Kudenov ?Laonice sp. Malacoceros sp. Polydora socialis (Schmarda) Polydora cf. woodwicki Polydora sp. Prionospio cirrifera Wiren Prionospio cf. cirrifera Prionospio multipinnulata Blake & Kudenov Spo pacifica Blake & Kudenov FAMILY: MAGELONIDAE SUB ORDER: CHAETOPTERIFORMIA FAMILY: CHAETOPTERIDAE Mesochaetopterus sp. SUB ORDER: CIRRATULIFORMIA FAMILY: CIRRATULIDAE Cirriformia capensis (Schmarda) Cirriformia filigera (delle Chiaje) Dodecaceria sp. ORDER: CAPITELLIDA FAMILY: CAPITELLIDAE Barantolla lepte Hutchings Capitella capitata’ (Fabricius) Letocapitella sp. Notomastus torquatus Hutchings & Rainer PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 FREQUENCY/ ABUNDANCY o/f u/a r/n u/a r/a r/a o/f o/f u/a o/f o/f u/f o/f a iilaee tiles ilans tiles iis ins ie ° =< eS o/f FOUND REGISTRATION AT SITES Nos 27 G15228 DB W201435-6 1,2,3,5,6,7 W201433-4 ahr) 0 W201431-2 2,3,9,/ W201429-30 2,6 W201427-8 4,7 W201426 7 W201425 1,2,3,4,5,6 W201397 1 W 201400 1 W201399 1,2,3,4,6 W201398 5 W202618 3,5 W201392 7 W 201396 Phsd)e7) W201391 2 W 201388 3 W201390 DoT W201393 6 W 201387 3,7 W 201389 5 W201394 3,5 W201385 2,3 W201395 45 W 201386 DDS) 0 W201384 6 W201383 5 W201382 7 W199764 73.7) W202616 2s) W202617 2 W199748 5 W201381 1 W20138 7 W201378 5 W201379 P. A. HUTCHINGS, J. T. VANDER VELDEANDS. J. KEABLE Appendix cont'd. FAMILY: MALDANIDAE Axiothella sp. Euclymene trinalis Hutchings ORDER: OPHELIDA FAMILY: OPHELIIDAE FAMILY: SCALIBREGMIDAE ORDER: PHYLLODOCIDA SUB ORDER: PHYLLODOCIFORMIA FAMILY: PHY _LLODOCIDAE Anaitides longipes (Kinberg) Eumida cf. sanguinea Phyllodoce novaehollandiae Kinberg Phyllodoce sp.1 Phyllodocid sp.1 Phyllodocid sp.2 Phyllodocid sp.3 SUB ORDER: APHRODITIFORMIA FAMILY: POLYNOIDAE Harmothoe sp.1 Harmothoe sp.2 Lepidasthenia sp. Lepidonotus carinulatus Grube Lepidonotus melanogrammus Haswell Lepidonotus n.sp.1 Lepidonotus n.sp.2 Lepidonotus n.sp.3 Lepidonotus n.sp.4 FAMILY: SIGALIONIDAE Psammolyce cf. antipoda Sigalion bandaeensis Horst FAMILY: CHRYSOPETALIDAE Chrysopetalum sp.1 SUB ORDER: NEREIDIFORMIA FAMILY: HESIONIDAE Podarke angustifrons (Grube) FAMILY: SYLLIDAE Autolytus sp.1 Autolytus sp.2 Autolytus sp.3 Autolytus sp.4 Syllid ?sp.1 Syllid ?sp.2 Syllid ?sp.3 Syllid ?sp.4 Syllid ?sp.5 Syllid ?sp.6 FAMILY: NEREIDIDAE Australonereis ehlersi (Augener) Ceratonereis aequisetis Augener Nereis maxillodentata Hutchings & Turvey Nereis cf. triangularis Perinereis amblyodonta (Schmarda) Platynereis dumerilit antipoda Hartman SUB ORDER: GLYCERIFORMIA FAMILY: GLYCERIDAE Glycera tridactyla Schmarda u/a o/f u/f o/f r/a r/a r/a u/f o/f o/f o/f u/f r/a o/f o/f o/f r/a r/a r/a o/f u/f o/f r/a r/a u/f o/f o/f o/f r/n r/n r/a o/f o/f u/f r/a r/n o/f o/f r/n o/f o/f 2a ONT NNN 2 1,2,3,5,6,7 LAG DIS] 4,7 2,7 DG) 1,2,5,6 1,2,3,4,5,6,7 7 7 1,2,3,4,5,6,7 Dba) 3)8) 351 W199732-41 W199742 W202619 W201412 W 202472 entire family currently being revised W 201418 W201417 W201422 W201421 W 201420 W 201416 W 201415 W 201414 W201419 W 201413 W199763 W199721-31 W 201374 W 201364 W 201365 W 201366 W 201367 W 201368 W 201369 W 201370 W 201371 W201372 W 201373 W 201361 W201362 W 202521 W202607 W 201363 W 202610 W199743-7 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 DoD BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix contd. FAMILY: NEPHTYIDAE Nephtys australiensis Fauchald ORDER: AMPHINOMIDA FAMILY: AMPHINOMIDAE Euphrosine n.sp. ORDER: EUNICIDA FAMILY: EUNICIDAE Eunice aphroditois (Pallas) Eunice cf. australis Eunice torrestensis McIntosh Eunice tridentata Ehlers Eunice tubifex Crossland Marphysa sanguinea (Montagu) Nematonereis unicornis (Grube) Falola sp.1 FAMILY: ONUPHIDAE Diopatra dentata Kinberg FAMILY: LUMBRINEREIDAE Augeneria verdis Hutchings & Murray Lumbineris latreilli Audouin & Milne Edwards FAMILY: ARABELLIDAE Arabella tricolor iricolor (Montagu) Arabella n.sp.1 FAMILY: LYSARETIDAE Lysidice cf. collaris Lysidice ninetta Audouin & Milne Edwards ?Lysidice sp. FAMILY: DORVILLEIDAE Dorvillea australiensis (McIntosh) Protodorvillea sp. Schistomeringos loveni (Kinberg) ORDER: OWENIIDA FAMILY: OWENIIDAE Owenia fusiformis delle Chiaje ORDER: TEREBELLIDA FAMILY: SABELLARIIDAE Idanthyrsus pennatus (Peters) FAMILY: AMPHARETIDAE Tsolda pulchella Muller FAMILY: TEREBELLIDAE Lanassa ocellata Hutchings & Glasby Lanice bidewa Hutchings & Glasby Longicarpus modesta Hutchings & Murray Nicolea amnis Hutchings & Murray Pista australis Hutchings & Glasby Pista violacea Hartmann-Schroder Reteterebella aloba Hutchings & Glasby Streblosoma acymatum Hutchings & Rainer Terebella pappus Hutchings & Murray Thelepus boja Hutchings & Glasby Thelepus brevicauda Hutchings & Glasby Thelepus extensus Hutchings & Glasby ORDER: SABELLIDA FAMILY: SABELLIDAE Amphiglena mediterranea (Leydig) Branchiomma nigromaculata (Baird) PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 r/n o/f r/a r/a o/f o/f o/f o/f u/f o/f r/a o/f o/f o/f r/n r/a o/f o/f o/f o/f u/f r/n u/a u/a o/f o/f u/a r/a u/f o/f u/f o/f u/a o/f o/f u/a u/f r/n 1 3)4259)-(6) 2,3,7 12 SE DROW 2,35 7 2 2p 6,7 2,3,5 13,45 950 2 Dah OM W200238-337 W199762 W 201411 W 201410 W 201409 W201408 W201407 W201406 W199804-5 W201405 W200191-202 W 201403 W 201402 W201401 W198896-976 W19773-802 W199803 W201404 W199751-4 W199749-50 W199755-9 W 200203-34 W199879-95 W199760-1 W 201375 W200388-97 W200375-80 W200635-7 W 200906 W200398 W 201377 200404-10 W198918 W 201376 W198916-7 W199827-30 W199831-70 P. A. HUTCHINGS, J. T. VANDER VELDEANDS. J. KEABLE Appendix contd. Megalomma sp. Sabellastarte indica (Savigny) FAMILY: SERPULIDAE Galeolaria caespitosa Savigny Hydroides cf. brachyacantha Neovermilia globula Dew Pomatoceros sp.1 Pomatoceros sp.2 Protula sp.1 Serpula jukesi Baird Serpula rubens Straughan Serpula sp.1 Spirobranchus tetraceros (Schmarda) PHYLUM: MOLLUSCA CLASS: POLYPLACOPHORA ORDER: NEOLORICATA FAMILY: LEPIDOPLEURIDAE Parachiton sp. FAMILY: ISCHNOCHITONIDAE Callistochiton antiquus (Reeve) Ischnochiton australis Sowerby Ischnochiton cariosus (Dall) Ischnochiton elongatus crispus Reeve Ischnochiton lentiginosus (Sowerby) Ischnochiton smaragdinus (Angas) Ischnochiton versicolor Iredale & Hull Ischnochiton sp.1 Ischnochiton sp.2 FAMILY: MOPALIIDAE Plaxiphora albida (Blainville) Plaxiphora matthews: Iredale FAMILY: SCHIZOCHITONIDAE Lorica volvox (Reeve) FAMILY: CHITONIDAE Chiton jugosus Gould Chiton pelliserpentis maugeanus (Iredale & May) FAMILY: ACANTHOCHITONIDAE Acanthochitona pilsbryi (Sykes) Acanthochitona retrojecta (Pilsbry) Acanthochitona sp.1 Notoplax cf. rubrostrata CLASS: BIVALVIA ORDER: NUCULOIDA FAMILY: NUCULANIDAE Nuculana spathula (Hedley) Scaeoleda hanley: (Angas) ORDER: SOLEMYOIDA FAMILY: SOLEMYIDAE Solemya australis (Lamarck) ORDER: ARCOIDA FAMILY: ARCIDAE Barbatia pistachia (Lamarck) Acar botanica (Hedley) FAMILY: GLYCY MERIDIDAE Glycymeris flammeus (Reeve) r/a u/f r/n r/a o/f r/a r/n o/f u/f u/a u/f o/f o/f o/f r/n r/n r/n u/f u/a r/a o/f o/f r/a u/f o/f r/f r/n r/n r/n u/f u/f o/f r/n u/a u/f o/f o/a 2,3 8,9 353 W199812-26 W199765-72 W 201358 W 201357 W 201355 W 201356 W 201360 W 201354 W 201359 W 201352 W 201353 W201351 C148803-4 C150544 C148821-3 C148813 C148824-5 C148820 C148814-6 C148817-9 C148826 C148827 C148802 C148807-8 C150545 C148797 C148798-800 C148809-10 C1488il C148812 C148805-6 C150568 C150570 C148522 C148542 C148541 C148794-5 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 354 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix cont'd. ORDER: MYTILOIDA FAMILY: MYTILIDAE Mytilus edulis (Linnaeus) Austromytilus rostratus Dunker Trichomya hirsuta (Lamarck) Musculus nanus (Dunker) Musculus sp.(juv.) Trichomusculus barbatus Reeve Modiolus areolatus (Gould) Xenostrobus securis (Lamarck) ORDER: PTERIOIDA FAMILY: PTERIIDAE Electroma georgiana (Quoy & Gaimard) FAMILY: PULVINITIDAE Vulsella vulsella (Linnaeus) FAMILY: PECTINIDAE Mimachlamys asperrimus (Lamarck) FAMILY: ANOMIIDAE Anomia descripta Iredale Anomia tone (Gray) Monia zealandica (Gray) FAMILY: LIMIDAE Lima nimbifer (Iredale) FAMILY: OSTREIDAE Ostraea angasi Sowerby Crassostrea gigas (Thunberg) Saccostrea commercialis (Iredale & Roughley) ORDER: VENEROIDA FAMILY: LUCINIDAE Wallucina assimilis (Angas) FAMILY: ERYCINIDAE Lasaea australis (Lamarck) FAMILY: GALEOMMATIDAE Ambuscintilla praemium Iredale FAMILY: CARDITIDAE Cardita excavata Deshayes FAMILY: CORDIIDAE Acrosterigma cygnorum (Deshayes) Fulvia tenuicostata (Lamarck) FAMILY: MACTRIDAE Spisula trigonella (Lamarck) FAMILY: MESODESMATIDAE Mesodesma elongata (Reeve) FAMILY: SOLENIDAE Solen vaginoides (Lamarck) FAMILY: TELLINIDAE Tellina deltoidalis (Lamarck) Tellina tenuslirata Sowerby Tellina sp. FAMILY: DONACIDAE Donax sp. (juv.) FAMILY: PSAMMOBIIDAE Gari sp. Sanguinolaria donacioides (Reeve) PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 r/n o/f r/n r/n u/n r/n u/f o/f o/f o/f u/f o/f u/f o/f o/f u/f o/f r/n u/f r/n r/a o/f o/f o/f o/f o/f o/f r/n u/f r/a o/f o/f o/f 2,3,5,6,7 7 2,5,7 7 Boll go) 7 7 2 Daal] D7) 5 2,5,6,7 2,5,6,7 1,3,4,5,6 9 3,9 5,6 C148548-9 C150549 C148538-40 C148533 C148534 C148537 C148532 C150577 C150558 C148535 C150563 C150548 C148530 C150564 C148536 C148793 C150556 C148796 C148523 C148545-6 C150547 C150553 C150546 C148521 C150572 C148516 C150571 C148517 C150573 C150574-5 C150557 C150559 C148543 P. A. HUTCHINGS, J. T. VANDER VELDEANDS. J. KEABLE Appendix contd. FAMILY: VENERIDAE ? Notocallista sp. Eumarcia fumigata (Sowerby) Venerupis crenata (Lamarck) Venerupis fabagella (Deshayes) Bassina pachyphylla (Jonas) Placamen placidum (Philippi) Timoclea cardiodes (Lamarck) ORDER: MYOIDA FAMILY: CORBULIDAE Corbula smithiana Brazier FAMILY: TEREDINIDAE Teredo sp. Bankia sp. FAMILY: HIATELLIDAE Fiatella australis (Lamarck) ORDER: PHOLADOMYOIDA FAMILY: LATERNULIDAE Laternula creccina Reeve Laternula cf. creccina FAMILY: MYOCHAMIDAE Myadora pandoriformis (Stutchbury) CLASS: GASTROPODA SUBCLASS: PROSOBRANCHIA ORDER: ARCHAEOGASTROPODA FAMILY: HALIOTIDAE Halvotis ruber (Leach) FAMILY: FISSURELLIDAE Amblychilepas javanicensis (Lamarck) Diodora lineata (Sowerby) Emarginula sp. Scutus antipodes Montfort Scutus sp. (juv.) FAMILY: PATELLIDAE Cellana tramoserica (Holten) ? Cellana tramoserica Patella chapmani (Tenison-Woods) Patella peroni (Blainville) FAMILY: ACMAEIDAE Notoacmea flammea (Quoy & Gaimard) Notoacmea petterdi (Tenison-Woods) Patelloida alticostata (Angas) Patelloida latistrigata (Angas) Patelloida mimula (Iredale) Patelloida mufria (Hedley) FAMILY: TROCHIDAE Austrocochlea concamerata (Wood) Austrocochlea constricta (Lamarck) Austrocochlea sp. (juv.) Bankivia fasciata (Menke) Cantharidella picturata (A. Adams & Angas) Euriclanculus floridus (Philippi) Eurytrochus stranger (A. Adams) Gena impertusa (Burrow) Granata imbricata (Lamarck) Herpetopoma aspersa (Philippi) o/f r/a r/n u/a o/f o/f o/f o/f o/f o/f r/n o/f o/f o/f u/f o/f o/f o/f r/a u/f r/n o/f u/f r/a u/f r/n r/n r/a r/n r/n r/n r/n o/f u/n u/a r/f r/n r/a r/f r/n 7 1;35455,6 2 NOWT 2,3,7 9 9 3 D9) 399 C150567 C148525 C148518 C148519-20 C150551 C150569 C148524 C150554-5 C150576 C150550 C148531 C150560-1 C148544 C150565 C148851 C150578 C150580 C148850 C148848 C148849 C148828-9 C150579 C148845-7 C148842-3 C148844 C150584 C148841 C148830-1 C148832-4 C148835-7 C148838-40 C148493 C148496-7 C148498 C148390 C148374-5 C148372-3 C150581 C148369 C148370-1 C148367-8 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 356 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix contd. Kerguelenella stoweai (Verco) o/f 2 C150582 Lewopyrga lineolaris (Gould) u/n 9 C148389 Mesoclanculus plebejus (Philippi) u/a 2 C150583 Montfortula rugosa (Quoy & Gaimard) r/n Zadl C148852 Phasianotrochus eximus (Perry) o/f 2 C148350 Phasianotrochus sp. u/a 2,3 C150585 Thaliota sp. u/f Dall C148357 Tugali sp. u/f Dell C148861 FAMILY: TURBINIDAE Astraea tenforiformis sirius (Gould) r/a 2,7 C148499 Astralium sp. o/f 2 C148500 Turbo torquatus (Gmelin) o/f 2,7 C148502 Turbo undulatus (Solander) r/a Doo) C148364-5 FAMILY: NERITIDAE Nerita atramentosa (Reeve) r/n DEO) C148494 ORDER: MESOGASTROPODA FAMILY: LITTORINIDAE Bembicium auratum (Quoy & Gaimard) r/n 22556 C148510-1 Bembicium nanum (Lamarck) r/n 5,7 C148508-9 Nodilittorina pyramidalis (Quoy & Gaimard) u/a 2,5,7 C148346 Nodilittorina unifasciata (Gray) r/n 2a OMT C148345 FAMILY: ASSIMINEIDAE Assiminea tasmanica Tenison-Woods o/f 1 C150586 FAMILY: ARCHITECTONICIDAE Philippia lutea (Lamarck; o/f 7 C148366 FAMILY: VERMETIDAE Serpulorbis stpho (Lamarck) o/f 7 C148507 FAMILY: PLANAXIDAE Hinea braziliana Lamarck o/a 27 C15088-9 FAMILY: POTAMIDIDAE Pyrazus ebeninus (Bruguiere) r/f 5,6 C150590 Velacumantus australis (Quoy & Gaimard) r/n 5,6 C148391-2 FAMILY: CERITHIIDAE Bittium sp. r/n Deka tl C148405-7 FAMILY: CREPIDULIDAE Crepidula aculeata Gmelin o/f 2 C150587 FAMILY: NATICIDAE Polinices melastomum (Swainson) o/f 8) C148354 FAMILY: RANELLIDAE Cabestana spengleri (Perry) o/f 8 C148503 Charonia lampax rubicunda (Perry) o/f 8 C148506 Ranella australasia (Perry) u/f “pil C148504-5 Sepla parthenopea (von Salis) o/f 7 C150591 ORDER: NEOGASTROPODA FAMILY: MURICIDAE Agnew2a tritoniformis (Blainville) r/a 2,7 C148358 Bedeva hanley: (Angas) r/a 2,3,5,6 C1548361 Bedeva sp. (juv.) o/f 3 C148362 Chicoreus denudatus (Perry) o/f 7 C148359 Haustrum vinosum (Quoy & Gaimard) r/a 2 C148355 ‘Lepsiella’ reticulata (Blainville) u/f 2 C148356 Mesoginella cf. translucida o/f 7 C148387 Mesoginella turbinata (Sowerby) u/a 9 C148388 Morula marginalba (Blainvilie) r/n 2 C148495 Thais orbita (Gmelin) r/n oll C148501 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. HUTCHINGS, J. T. VAN DER VELDEANDS. J. KEABLE Appendix contd. FAMILY: COLUMBELLIDAE Mitrella leucostoma (Gaskoin) Mitrella lincolnensis (Reeve) Miutrella pulla (Gaskoin) Mitrella sp. FAMILY: BUCCINIDAE Cominella lineolata (Lamarck) FAMILY: NASSARITDAE Nassarius burchardi (Dunker) Nassarwus cf. burchardi Nassarius glans particeps (Hedley) Nassarius pauperatus (Lamarck) Nassarius pauperus (Gould) Nassarius sp. (juv.) FAMILY: OLIVIDAE Olivella leucozona Adams & Angas FAMILY: TURRIDAE Guraleus pictus (Adams & Angas) FAMILY: CONIDAE Conus anemone Lamarck SUBCLASS: OPISTHOBRANCHIA ORDER: BULLOMORPHA FAMILY: BULLIDAE Bulla quoyii (Gray) FAMILY: PHILINIDAE Philine cf. angasi ORDER: ANASPIDEA FAMILY: APLYSIDAE Aplysia sydneyensis Sowerby ORDER: NOTOSPIDEA FAMILY: PLEUROBRANCHIDAE Pleurobranch sp. 1 ORDER: NUDIBRANCHIA SUBORDER: DORIDACEA FAMILY: DORIDIDAE Aphelodoris varia (Abraham) Dorid Nudibranch sp. 1 Dorid Nudibranch sp. 2 FAMILY: CHROMODORIDIDAE Ceratosoma amoena (Cheeseman) FAMILY: POLYCERIDAE Polycera capensis (Quoy & Gaimard) SUBORDER: AEOLIDACEA FAMILY: GLAUCIDAE Austraeolis cacaotica (Stimpson) Pteraeolidia tanthina (Angas) SUBORDER: ARMINACEA FAMILY: JANOLIDAE Caldukia affinis (Burn) SUBCLASS: PULMONATA ORDER: SYSTELLOMMATOPHORA FAMILY: ELLOBIIDAE Ophicardelus ornatus (Ferussac) Ophicardelus quoyi (H. & A. Adams) o/f o/f u/f u/f r/a r/a o/f o/f r/n o/f u/a o/f u/f o/f o/f o/f o/f o/f o/f o/f o/f o/f o/f o/f o/f o/f r/n r/a “I D 357 C148377-8 C148379-80 C148381-3 C148384-6 C148352-3 C148402 C148401 C148403 C148400 C148404 C148399 C148376 C148363 C148351 C148360 C148866 C148512 C148865 C150593 C148862 C148863 C150596 C150597 C150594 C150598 C150595 C148393-4 C148396-7 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 358 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix Cont'd. Ophicardelus cf. quoyt Ophicardelus sulcatus (H. & A. Adams) FAMILY: ONCHIDIIDAE Onchidella patelloides (Quoy & Gaimard) Onchidina australis (Semper) ORDER: BASOMMATOPHORA FAMILY: SIPHONARIIDAE Salinator fragilis (Lamarck) Salinator solida (von Martens) Siphonaria denticulata Quoy & Gaimard Stphonaria diemenensis Quoy & Gaimard Stphonaria funiculata Reeve Stphonaria sp. (juv.) ORDER: SIGMURETHRA Theba pisana (Muller) CLASS: SCAPHOPODA ORDER: GADILIDA FAMILY: SIPHONODENTALIIDAE Cadulus acuminatus Tate PHYLUM: ARTHROPODA CLASS CRUSTACEA SUBCLASS: OSTRACODA ORDER: MYODOCOPA Myodocopa spp. SUBCLASS: CIRRIPEDIA ORDER: THORACICA Austrobalanus imperator (Darwin) Austromegabalanus nigrescens (Lamarck) Balanus trigonus Darwin Balanus variegatus Darwin Catomerus polymerus (Darwin) Chamaesipho columna (Spengler) Chthamalus antennatus Darwin Elminius covertus Foster Ibla quadrivalvis Cuvier Notomegabalanus algicola (Pilsbry) Tesseropora rosea Krauss Tetraclitella purpurascens (Wood) SUBCLASS: MALACOSTRACA ORDER: LEPTOSTRACA Neballiacean sp. SUPERORDER: PERACARIDA ORDER: MYSIDACEA Hetzromysis sp. Striella australis W. M. Tattersall ORDER: CUMACEA Cumacean spp. ORDER: TANAIDACEA Tanaidacean spp. ORDER: ISOPODA SUBORDER: GNATHIIDEA Gnathia ferox (Haswell) SUBORDER: ANTHURIDEA Apanthura drosera Poore & Lew Ton PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 o/f u/f r/a r/f r/n r/n r/a r/n r/a o/f o/f u/n r/n a Wt sent ets Moo ey tea o/f u/f o/f r/n r/n u/f o/f 2,3,7,9 1,3,5,7,9 2,3,7,9 By/, C148395 C148398 C148859-60 C148864 C148347-8 C148349 C148853 C148854-5 C148856 C148857-8 C150600 C1i50969 P36562-83 P36114 P36119 P36112 P36614 P36122 P36120 P36124 P36125 P36113 P36116-8 P36121 P36123 P36584-7 P36617-9 P36615-6 P36594-613 P36525-48 P36588-91 P36050 P. A. HUTCHINGS, J. T. VAN DER VELDEANDS. J. KEABLE Appendix contd. Apanthura isotoma Poore & Lew ‘Ton Apanthura xanthorrhoea Poore & Lew Ton Apanthuretta olearia Poore & Lew Ton Bullowanthura pambula Poore Cyathura hakea Poore & Lew Ton Haliophasma canale Poore Haliophasma sp. 1 Haliophasma sp. 2 Leptanthura diemenensis (Haswell) Mesanthura dianella Poore & Lew Ton Paranthura acacia Poore Paranthura senecio Poore Ulakanthura marlee Poore SUBORDER: FLABELLIFERA Amphoroidea angustata Baker Cercets ?obtusa Cirolana australiense Hale Cirolana victoriae Bruce Cilicaea tenuicauda Haswell Ciltcaeopsis cf. whiteleggei Cilicaeopsis sp. Cymodoce cf. bidentata Cymodoce haswelli Harrison & Holdich Cymodoce spp. (females) ? Cymodopsis sp. Cymothoid sp. Eurydice ?binda Eurydice sp. Eurylana arcuata (Hale) Exosphaeroma sp. Haswellia carnea (Haswell) Haswellia cf. juxtacarnea Ischromene ? polytyla Limnoria sp. ? Paracilicaea sp. Paracassidina pectinata Baker Pseudolana towrae Bruce Serolis minuta Beddard Sphaeromatid sp. 1 u/f r/a o/f o/f o/f o/f o/f o/f r/a u/f o/f r/f o/f o/f o/f r/n o/f o/f o/f o/f o/f u/f r/n o/f o/f o/f o/f r/f o/a o/f o/f r/n r/a u/f o/f o/f o/f o/f 359 P36046-8 P35640-2 P36154-6 P36049 P36059 P36052 P36593 P36058 P36061-2 P36592 P36056-7 P36133-5 P35648-9 P36053-5 P36170-2 P35643-5 P36149 P36060 P35646-7 P36167-9 P36051 P35957 P36625 P35961 P36146-8 P35972 P36626 P36627 P35970 P35969 P36178 P36628 P35948-9 P35954 P35960 P36145 P36173-4 P36623-4 P35955 P36631 P36629 P35971 P35958 P36175-7 P35959 P35950 P36144 P35951 P36159-60 P35953 P36165-6 P35952 P36157-8 P36622 P35963 P36179 P35966-7 P35964-5 P35956 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 360 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix cont'd. Sphaeromatid sp. 2 Sphaeromatid sp. 3 Sphaeromatid sp. 4 Sphaeromatid sp. 5 Syncassidina aesturia Baker SUBORDER: ONISCOIDEA Oniscoidean sp. 1 Oniscoidean sp. 2 SUBORDER: VALVIFERA Euidotea cf. peronit Microarcturus sp. Neoarcturus sp. Pseudarcturella sp. Synidotea sp. SUBORDER: ASELLOTA Taniropsis sp. Tathrippa sp. Jaeropsis sp. 1 Jaeropsis sp. 2 Janirid sp. 1 Janirid sp. 2 Stenetrium armatum Haswell Stenetrium cf. armatum ORDER: AMPHIPODA SUBORDER: GAMMARIDEA FAMILY: AMPELISCIDAE Ampelisca dimboola Lowry & Poore Ampelisca euroa Lowry & Poore Byblis bega Lowry & Poore FAMILY: AMPHILOCHIDAE Cyproidea ornata Haswell Narapheonoides mullaya J. L.Barnard FAMILY: AMPITHOIDAE Ampithoe sp. 1 Ampithoe sp. 2 Ampithoe sp. 3 Ampithoe sp. 4 Ampithoe sp. 5 Cymadusa sp. 1 Cymadusa sp. 2 ? Pseudopleonexes sp. FAMILY: ANAMIXIDAE Anamixis sp. FAMILY: AORIDAE Aora hebes Myers & Moore Aora maculata (Thomson) Aora mortont (Haswell) Lemboides australis (Haswell) PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 o/f u/f o/f o/f u/f o/f o/f o/f o/f o/f o/f o/f r/n r/n r/n o/f o/f o/f u/n o/f o/f r/f o/f r/n o/f o/a u/f u/f o/f o/f u/f o/f o/f o/f r/n u/n r/n r/a wownonwnowo vo Kec) ~N NWnN™NNM ND PO P35962 P36620 P36621 P36630 P35968 P36143 P36633-6 P36632 P36068 P36071 P36070 P36072 P36069 P35658-9 P36152-3 P35653-5 P36150-1 P35660-2 P36163-4 P36064-5 P36066 P36067 P35650-2 P36161-2 P36063 P36660 P36638 P36659 P36709 P36708 P36005 P36712 P36706 P36713 P36707 P35993 P36710 P36705 P36651 P36714 P35992 P36187-8 P36014-5 P36032 P36181 P36031 P36189 P. A. HUTCHINGS, J. T. VAN DER VELDE AND S. J. KEABLE Appendix cont'd. Lembos aequimanus Schellenberg Xenocherra fasciata Haswell FAMILY: COROPHIIDAE Corophium sp. Paracorophium Pexcavatum Siphonoecetes spp. FAMILY DEXAMINIDAE Atylus homochir Haswell Haustoriopsis sp. 1 Haustoriopsis sp. 2 Paradexamine churinga J. L. Barnard Paradexamine dandaloo J. L. Barnard Paradexamine frinsdorfi Sheard Paradexamine lanacoura J. L. Barnard Paradexamine ?quarallia Paradexamine ? thadalee Polychetra tenuipes Haswell Syndexamine cf. runde Syndexamine sp. 1 FAMILY: EUSIRIDAE Gondogeneia microdeuteropa (Haswell) Meteusiroides sp. ? Paramoera sp. Tethygeneia nalgo J. L. Barnard Tethygeneia waminda J. L. Barnard FAMILY: EXOEDICEROTIDAE Exoediceroides maculosus (Sheard) FAMILY: HYALIDAE Hyale crassicornis (Haswell) Hyale cf. loorea Hyale maroubrae Stebbing Hyale rubra (Thomson) Hyale sp. FAMILY: ISAEIDAE Ampelisciphotis sp. Cheiriphotis sp. Gammaropsis sp. ?Gammaropsis sp. 1 ?Gammaropsts sp. 2 ?Gammaropsis sp. 3 Photis sp. 1 Photis sp. 2 FAMILY: ISCHYROCERIIDAE Cerapus sp. Ericthonius sp. Parajassa sp. FAMILY: LEUCOTHOIDAE Leucothoe assimilis J. L. Barnard r/f o/f r/n r/n o/f r/a r/a o/f o/f o/f u/f r/a o/f r/n o/f o/f u/f r/a r/a o/f r/n o/n o/f r/f u/a u/f u/f r/f r/n o/f r/a r/n u/f r/a r/n o/f o/f r/n u/n o/f NOWMN OND WW PD 1S<) 1S%) ~s No™wN 2,7 BeSol) Dro 8) 2,3,7 2,7 ll Dell eS EO 9 361 P36033 P36180 P36007 P36641 P36042 P36295-306 P36029 P36028 P36027 P36005 P36043 P35991 P36026 P36711 P36025 P36203 P36703 P36009 P36024 P35973 P36004 P36012 P36010 P36040 P35639 P35977 P36185-6 P36645 P36013 P36184 P36182-3 P35976 P36646 P36030 P35994-5 P36003 P35985 P36002 P35984 P36034-5 P36190-1 P36018 P36102-4 P36195-202 P36006 P36023 P36704 P36642 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 362 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix contd. Leucothoe boolpooli J. L. Barnard Leucothoe commensalis Haswell Paraleucothoe novaehollandiae (Haswell) FAMILY: LILJEBORGIIDAE Liljeborgia aequabilis Stebbing Liljeborgia sp. 1 Liljeborgia sp. 2 FAMILY: LYSIANASSIDAE Amaryllis sp. 1 Amaryllis sp. 2 Amaryllis sp. 3 Lysianassid sp. 1 Parawaldeckia dilkera J. L. Barnard Parawaldeckia sp. Tryphosella camelus (Stebbing) Uristidid sp. 1 Waldeckia sp. FAMILY: MELITIDAE Ceradocus ramsayi (Haswell) Ceradocus rubromaculatus (Stimpson) Ceradocus serratus (Bate) Ceradocus sp. ? Ceradocus sp. Dulichielia australis (Haswell) Elasmopus bollonsi Chilton Elasmopus ?yunde Gammarella berringar (J. L. Barnard) Gammarella mokari (J. L. Barnard) Maera viridis Haswell Maera sp. ?Maera sp. Mallacoota subcarinata Haswell Melita matilda J. L. Barnard Melita sp. Victoriopisa australiensis (Chilton) FAMILY: PARACALLIOPIIDAE Paracalliope sp. FAMILY: PHLIANTIDAE Gabophtias olono J. L. Barnard [phiplatera whitelegger Stebbing PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 r/n r/a r/a r/a o/f o/f o/f o/f r/a u/f r/f r/f u/f o/f r/n r/n r/f u/f o/f r/f r/n u/n o/f o/f u/f r/n o/f r/n o/f r/a u/f r/a r/n u/f o/f oo dnll eho l) Dell 2,3,7 P35990 P36213 P35983 P36214 P35989 P36215 P36001 P36216 P36022 P36038 P36700 P36701 P35982 P36702 P35981 P36212 P36021 P36211 P36207-9 P36044-5 P36016-7 P36210 P36204-6 P36036 P35988 P35987 P36229 P36637 P36000 P36218-9 P36650 P36644 P35980 P36221 P35979 P36230 P36644 P35999 P35998 P36224-6 P35978 P36222-3 P36649 P36643 P35988 P36041 P36227-8 P36647-8 P36699 P35637-8 P36011 P36220 P36020 P. A. HUTCHINGS, J. T. VAN DER VELDE ANDS. J. KEABLE Appendix cont'd. FAMILY: PHOXOCEPHALIDAE Birubius mayamayi Barnard & Drummond Birubius muldarpus Barnard & Drummond Birubius cf. nammuldus Birubius quearus Barnard & Drummond ?Burubius sp. Brolgus tattersalli (J. L. Barnard) Tipimegus dinjerrus Barnard & Drummond Wildus sp. FAMILY: PODOCERIDAE Podocerus sp. 1 Podocerus sp. 2 Podocerus sp. 3 FAMILY: SYNOPIIDAE Tiron sp. FAMILY: TALITRIDAE Orchestia ?australts Orchestia sp. FAMILY: UROHAUSTORIIDAE Gheegerus cf. garbaius Tottungus ?tungus Urohaustorius gunni Barnard & Drummond Urohaustorius merkanius Barnard & Drummond Urohaustorius metungi Fearn-Wannan Urohaustorius parnggius Barnard & Drummond Urohaustorius ?urungari SUBORDER: CAPRELLIDEA Caprella danilevskit Czerniavski Caprella equilibra Say Caprella scaura Templeton ? Hircella cornigera Orthoprotella cf. mayert Paraproto spinosa (Haswell) ? Paraproto sp. SUPERORDER: EUCARIDA ORDER: DECAPODA SUBORDER: NATANTIA SECTION: PENAEIDEA Penaeus plebejus Hess ? Penaeus sp. (juv.) SECTION: CARIDEA Alpheus euphrosyne richardsoni Yaldwyn Alpheus socialis Heller Alpheus sp. Alope ?ortentalis Hippolyte caradina Holthius Hippolyte ventricosa Milne-Edwards Palaemon affinis Milne-Edwards Palaemon serenus Heller Rhynchocinetes ?rugulosus Synalpheus tumidomanus (Paulson) o/f o/f o/f r/f o/f o/f o/f r/a o/a r/n r/a o/f r/f u/f o/f o/f o/f o/f u/f o/f o/f o/f o/f o/f u/f o/a u/f u/f o/f u/f u/f u/f olf u/a o/f r/a o/f o/a u/f WWUONUOUONN™N DY 1S) woVmonwVnowonwo 1,3,4,5,6 363 P36691 P36698 P36690 P36694-5 P36697 P36692 P36696 P36693 P35997 P35996 P36232-3 P36019 P36231 P36039 P36037 P36217 P35974 P35975 P36661 P36652 P36658 P36656 P36653-4 P36657 P36655 P36684-5 P36688-9 P36676-7 P36687 P36681-3 P36686 P36678-90 P36102-3 P36668 P36106 P36100 P36666 P36115 P36664-5 P36663 P36108 P36110 P36101 P36667 P36662 P36105 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 364 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix contd. SUBORDER: REPTANTIA SECTION: MACRURA SUPERFAMILY: THALASSINIDEA Axiopsis australiensis De Man Callianassa arenosa Poore SECTION: ANOMURA Diogenes custos affinis Henderson Diogenes senex Heller Paguristes squamosus McCulloch Pagurus ?lacertosus nana Pagurus sinuatus (Stimpson) Pagurus sp. SECTION: BRACHYURA Actaea peronit (Milne-Edwards) Amarinus paralacustris (Lucas) Carcinus maenas (Linnaeus) Cyclograpsus audouinit Milne-Edwards Cyclograpsus cf. audouinit Halicarcinus ovatus Stimpson Heloecius cordiformis Milne-Edwards Helograpsus haswellianus (Whitelegge) Hymenosoma hodgkini Lucas Leptograpsus variegatus (Fabricius) Macrophthalmus latifrons Haswell Mictyris longicarpus Latreille Naxia deflexifrons Haswell Notomithrax minor (Filhol) Notomithrax ursus (Herbst) Ovalipes australiensis Stephenson & Rees Ovalipes sp. (juv.) Ozius truncatus (Milne-Edwards) Pachygrapsus laevimanus Stimpson Paragrapsus laevis (Dana) Petalomera lateralis (Gray) Pilumnus rufopunctatus Stimpson Pilumnus serratifrons (Kinahan) Pilumnus sp. Pinnotheres hickmani (Guiler) Plagusia chabrus (Linnaeus) Plagusia glabra Dana Portunus pelagicus (Linnaeus) Sesarma erythrodactyla Hess Xanthias elegans (Stimpson) ?Xanthid (juv.) Decapod Larvae CLASS: PYCNOGONIDA Pycnogonid spp. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 o/f u/f o/f o/f o/f o/f o/f o/f o/f u/f r/a r/n o/f r/n o/f r/n o/f r/a u/a r/n o/f u/a o/f o/f o/f r/a u/a r/n o/f r/a r/f o/f o/f r/f u/f o/f r/f o/f o/f o/f u/f NNN WW OO P36104 P36099 P36111 P36098 P36107 P36109 P36669 P36670 P36083 P36090 P36248-9 P36089 P36076 P36239 P36080 P36095 P36234-6 P36086 P36247 P36091 P36242-3 P36087 P36079 P36085 P36245-6 P36084 P36673 P36094 P36675 P36088 P36126 P36097 P36081 P36238 P36075 P36092 P36240-1 P36073 P36082 P36237 P36077 P36671 P36672 P36074 P36078 P36096 P36244 P36674 P36549-61 P. A. HUTCHINGS, J. T. VAN DER VELDE AND S. J. KEABLE Appendix cont'd. PHYLUM: SIPUNCULIDA Sipunculan spp. PHYLUM: BRACHIOPODA Magellania flavescens (Lamarck) PHYLUM: ECHINODERMATA CLASS: ASTEROIDEA ORDER: VALVATIDA FAMILY: ASTERINIDAE Patiriella calcar (Lamarck) Patiriella exigua (Lamarck) Patiriella gunni (Gray) FAMILY: OREASTERIDAE Nectria ocellata E. Perrier FAMILY: ASTEROPSEIDAE Petricia vernicina (Lamarck) ORDER: SPINULOSIDA FAMILY: ECHINASTERIDAE Plectaster decanus (Muller & Troschel) ORDER: FORCIPULATIDA FAMILY: ASTERIIDAE Coscinastertas calamaria (Gray) Uniophora granifera (Lamarck) CLASS: OPHIUROIDEA ORDER: OPHIURIDAE FAMILY: OPHIOCOMIDAE Clarkcoma pulchra (H. L. Clark) FAMILY: OPHIONEREIDAE Ophionerets schayert (Muller & Troschel) FAMILY: OPHIACANTHIDAE ? Ophiacantha sp. FAMILY: OPHIACTIDAE Ophiactis resiliens Lyman FAMILY: AMPHIRORIDAE Amphipholis squamata (Delle Chiaje) Amphuura constricta Lyman Amphiura micra H. L.Clark FAMILY: OPHIOTHRICIDAE. Ophiothrix (Ophiothrix) caespitosa Lyman Ophiothrix (Ophiothrix) cilaris (Lamarck) Ophiothrix (Placophiothrix) spongicola Stimpson CLASS: ECHINOIDEA ORDER: CIDAROIDA FAMILY: CIDARIDAE Phyllacanthus parvispinus Tenison-Woods r/a o/f u/a r/n u/a o/f u/a o/f r/n o/a o/f u/n o/f o/a of u/f o/f o/f o/f o/f r/f DES SONI Zell Dyn) 23h Zool 7 365 W 201423 C150562 Ji9849-52 Ji9887 Ji9865-73 J19875-76 J198788-92 J19853-54 J19874 J19855 J19856-8 Ji9864 J19884-5 J19834-48 J19859-63 Ji9832 J19826-9 J19895 J19902-5 Ji0008-9 J20002 J20001 J19899-901 J19881-3 Ji9906 J19877-79 Ji9907 J19893-4 J19995-7 Ji9831 Ji19823-5 PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 366 BENTHIC MACROFAUNA OF TWOFOLD BAY Appendix cont'd. ORDER: DIADEMATOIDA FAMILY: DIADEMATIDAE Centrostephanus rodgersti (A. Agassiz) ORDER: TEMNOPLEUROIDA FAMILY: TEMNOPLEURIDAE Holopneustes inflatus A. Agassiz ORDER: ECHINOIDA FAMILY: ECHINOMETRIDAE Heliocidaris erythrogramma (Valenciennes) Heliocidarts tuberculata (Lamarck) CLASS: HOLOTHUROIDEA ORDER: DENDROCHIROTIDA FAMILY: CUCUMARIIDAE Pentacta ignava (Ludwig) Pseudocnus sp. ORDER: MOLPAPIIDA FAMILY: CAUDINIDAE Paracaudina chilensis var. ransonnett: (von Marenzeller) CLASS: CRINOIDEA ORDER: COMATULIDA FAMILY: COMASTERIDAE Cenolta tasmaniae (A. H. Clark) PHYLUM: CHORDATA SUBPHYLUM: TUNICATA CLASS: ASCIDIACEA ORDER: PLEUROGONA FAMILY: STYELIDAE Styela plicata (Lesueur) FAMILY: PYURIDAE Herdmania momus (Savigny) Pyura gibbosa (Heller) Pyura spinifera (Quoy & Gaimard) Pyura stolonifera (Heller) PHYLUM: CHORDATA GRADE: PISCES ORDER: AMPHIOXIFORMES FAMILY: BRANCHIOSTOMATIDAE Amphioxus sp. ORDER: ANGUILLIFORMES FAMILY: ANGUILLIDAE Anguilla australis Richardson FAMILY: OPHICHTHIDAE Muraenichthys sp. ORDER: SCOR PAENIFORMES FAMILY: SCOR PAENIDAE Ruboralga ergastulorum (Richardson) Scorpaenidae sp. Centropogon australis {White) ORDER: PERCIFORMES FAMILY: CLINIDAE Heteroclinus heptaeolus (Ogilby) PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 r/a o/f r/a o/f u/a o/f o/f o/f r/n r/n o/f r/n o/f o/f o/f o/f o/f o/f o/f 2a, all 4,6 J19815 J19910 J19816-21 Ji9822 J19896-8 J19909 J19908 J19833 Y 2060 Y 2061-65 Y¥ 2057-59 Y 2049 Y 2050-56 126021-002 126017-001 126018-001 126019-001 126007-001 126015-003 126015-002 126008-001 126010-001 P. A. HUTCHINGS, J. T. VAN DER VELDE ANDS. J. KEABLE 367 Appendix cont'd. FAMILY: GOBIIDAE Arenigobius bifrenatus (Kner) o/f 5 126020-001 Favonigobius tamarensis (Johnson) o/f 5 126014-002 Philypnodon granaiceps (Kreft) o/f 126016-001 Pseudogobius olorum (Sauvage) o/f 1,5,6 126013-001 126014-001 126015-001 Pseudogobius sp. o/f 1 126011-001 126009-001 FAMILY: SERRANIDAE Epinephelus sp. o/f 7 126012-001 PROC, LINN. SOC. N.S.W., 110 (4), (1988) 1989 ear et oh eo ; aA ule Tats sett vie Hn : WARIS bat ssee iis a VP itee 0,6.) Se eter a art ‘ mn . Aeros fil » Ages “geenC LIS . Hi TOES ee) Peivigh ers) pesadtud dint) Waders eta li5 ‘erick boeing Cees. era APU lhe a eS i*) > ; ioc 0 nL “AAG fi uy a : i ) + wD} ‘ pare ' ri y y ry . vo oe ieee 5 , , (Aare trus Eames | & fies Be ee ‘ + = f iri, OF } 7 TAL iT, : 4 be % hae + bite = i¢ ' s ¢ i's ci SPe { oJ al [Ww j “4 04. *%e F Ir EO Sra ia) a a j 7 saree i) ree) Pi car | S4Reits . ths efi ely att 7 Ni . ‘ : 4 i i - t of i } The Dictyotales (Algae: Phaeophyta) of New South Wales P. A. FARRANT and R. J. KING FARRANT, P. A., KING, R. J. The Dictyotales (Algae: Phaeophyta) of New South Wales. Proc. Linn. Soc. N.S.W. 110(4), (1988) 1989: 369-405. A survey is given of the Dictyotales (Phaeophyta) on the New South Wales main- land coast. Twenty-two species are recognized in 13 genera: Duictyopteris, Dictyota, Dilophus, Distromium, Homoeostrichus, Lobophora, Lobospira, Pachydictyon, Padina, Spatoglos- sum, Stypopodium, Taonia, and Zonarza. For each, a concise description with illustrations, based on New South Wales material, is presented. Keys are provided to genera and species. P. A. Farrant, Royal Botanic Gardens Sydney, Mrs Macquarie’s Rd, Sydney, Australia 2000, and R. J. King, School of Biological Science, University of New South Wales, PO. Box 1 Kensington, Australia 2033, manuscript received 15 March 1988, accepted for publication 20 July 1988. INTRODUCTION Brown algae in the Order Dictyotales Kjellman are a conspicuous component of the lower eulittoral and sublittoral marine flora of the New South Wales coast (King and Farrant, 1987). Many specimens, especially immature plants, of the Dictyotales on the New South Wales coast have been difficult to identify despite publications from southern Queensland (Cribb, 1954), southern Australia (Womersley, 1967, 1987) and Lord Howe Island, New South Wales (31°33’S, 159°05’E) (Allender and Kraft, 1983). Lord Howe Island is administratively part of New South Wales, but it is as far removed from the mainland New South Wales coast as are much of Queensland and Victoria (Fig. 1). The Dictyotales is a distinctive order and members can be recognized by a flattened thallus with apical growth, tufts of surface hairs and oogamous reproduction with characteristic antheridia and oogonia on or embedded in the thallus surface (Womersley, 1987). The plants are usually erect or occasionally decumbent in habit. The fronds are either dichotomous, subdichotomous or irregular if growth is initiated by a single apical cell or a small group of apical cells, or flabellate or irregular if growth is from a row of marginal cells. The Dictyotales have been placed traditionally in a single family, the Dictyotaceae, within which two tribes were recognized: the Dictyoteae, characterized by a single apical cell, and the Zonarieae with a group or marginal row of apical cells. Womersley (1987) notes that two families, the Dictyotaceae and the Zonariaceae, may seem warranted, but that their recognition is not satisfactory for some juvenile stages. There are two genera which, though clearly members of the Dictyotales, are sufficiently distinct to warrant placement in separate families. Allender (1980) placed the genus Dictyotopsis ‘Troll in a new family, Dictyotopsidaceae. In this family the single apical cell segments laterally and the thallus is monostromatic. In those members of the Dictyotaceae which possess a single apical cell it is oriented transversely to the branch apex or is conical and new cells are cut off only from the lower side (Womersley, 1987). Dictyotopsis, represented by the single species D. propagulifera Troll, has not been found on New South Wales shores though sterile plants have been recorded by one of us (R.J.K.) associated with man- groves in Queensland (near the mouth of the Endeavour River and at Hinchinbrook Island). Womersley (1987) has described a new family, the Scoresbyellaceae, for the dis- tinctive monotypic genus Scoresbyella. The plants resemble those in the Dictyoteae, but PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 370 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Northern , Territory ; Pacific ; Ocean Queensland Western Australia South Australia Hastings Pt Ballina New South Wales , /Port Stephens Island § Sydney Jervis Bay Fig. 1. Locality map. the apical cell is oriented longitudinally and cells are cut off laterally from two faces rather than from one basal side. Scoresbyella profunda Womersley appears to be a rare alga and is thus far known from only a few deep-water collections in South Australia. The total number of genera in the order Dictyotales is 18 and there are approxi- mately 100 species. All genera except Stoechospermum have been recorded in Australia. Fifteen genera and 41 species occur on temperate southern Australian coasts (Womersley, 1987). This contrasts with the generalization that the order is best de- veloped on tropical and subtropical coasts (Haupt, 1932; Bold and Wynne, 1985). Table 1 shows the number of genera and species in various regions of Australia and indicates that the order Dictyotales is well represented in both northern and southern Australia. The high figure for southern Australia reflects the richness of the well-documented algal flora (Womersley, 1981, 1987). The comparable figure for northern Australia is based upon species records only (not including those for Lord Howe Island) (Lewis, 1985), and may be artificially high, especially for Dictyota. Of the 44 species recorded in northern Australia, approximately 24 are recorded for Queensland and Norfolk Island, a number similar to that for southern Queensland (20), New South Wales (22), and Lord Howe Island (22). MATERIALS AND METHODS Members of the Order Dictyotales were collected on a regular basis at Fairlight in Sydney Harbour (33°50’S, 151°15’E) in an associated study of the phenology of PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P.A. FARRANT ANDR. J. KING 371 TABLE 1 Number of species in the Order Dictyotales recorded for various regions of Australia: northern Australia (after Lewis, 1985), southern Queensland (after Lewis, 1985: Cribb, 1954; N.S.W. herbarium specimens), Lord Howe Island (Allender and Kraft, 1983), New South Wales (this study) and southern Australia (Womersley, 1987) nthn sthn Lord N.S.W. sthn Aust. Qld Howe Aust. Island ORDER DICTYOTALES F. DICTYOTACEAE Dictyoteae Dictyota 15 6 3 3 7 Dilophus 3 2 2 2 7 Glossophora 2 0 0 0 1 Pachydictyon 0 0 1 1 2 Zonarieae Chlanidophora 0 0 0 0 1 Dictyopteris 7 3 5 1 5 Distromium 0 0 1 1 2 Homoeostrichus 0 0 0 2 3 Lobophora 1 1 1 1 1 Lobospira 0 0 0 1 1 Padina 6 6 4 4 4 Spatoglossum 3 1 1 1 1 Stypopodium 3 0 2 1 0 Taonia 1 0 1 1 1 Zonaria 2 1 1 3 4 F. DICTYOTOPSIDACEAE Dictyotopsis 1 0 0 0 0 F. SCORESBYELLACEAE Scoresbyella 0 0 0 0 1 Total Genera: 17 Species: 44 20 22 22 41 members of the order (King and Farrant, 1987). In addition, collections were made at other localities in New South Wales, but the majority of collections made were from the Sydney region. Localities in the Sydney metropolitan area include Camp Cove, Dobroyd, Fairlight, Lady Jane Beach, Mrs Macquarie’s Point, Point Piper and Vaucluse in Sydney Harbour; Bare Island in Botany Bay; Church Point in Pittwater; and Boat Harbour, Clovelly, Collaroy, Fairy Bower, Harbord, Long Bay, Long Reef and Newport on the nearby open coast. Localities in the Jervis Bay area include Plantation Point (in- side the bay), Crookhaven Heads (north of the bay) and Steamers Beach (south of the bay). All New South Wales localities other than these can be found in the Reader’s Digest Atlas of Australia (1977) or in Fig. 1. Voucher specimens (herbarium sheets and wet material) are lodged in the John T. Waterhouse Herbarium, University of New South Wales (UNSW). Herbarium material from NSW (including the A. H. S. Lucas collection), UNSW, MELU, and selected specimens from MEL and MUCV were examined during the course of the study. RESULTS The order Dictyotales in New South Wales is represented by 22 species in 13 genera (Table 2). A key to the genera is presented below and this is followed by a description of each species based on mature specimens from the New South Wales coast. Where a PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 372 THE DICTYOTALES{ALGAE PHAEOPHYTA) OF NEW SOUTH WALES genus Is represented by more than one species, a key to the species is given. The descrip- tions are concise and do not repeat detailed information and complete lists of synonymy readily available elsewhere (especially in Allender and Kraft, 1983; and Womersley, 1987). Rather they concentrate on information necessary for the identification of the taxa. Only where a species has been widely referred to by another name is the synonym given. Seasonality, seasonal and spatial variation in abundance, and seasonal variation in fertility for some of the common species are given by King and Farrant (1987). TABLE 2 Species of Dictyotales in New South Wales Dictyopteris acrostichoides (J. Agardh) Boergesen Dictyota alternifida J. Agardh Dictyota bartayresit: Lamouroux Dictyota dichotoma (Hudson) Lamouroux Dilophus intermedius (Zanardini) Allender and Kraft Dilophus marginatus J. Agardh Distromium flabellatum Womersley Homoeostrichus olsenii Womersley Homoeostrichus sinclairit (Hooker and Harvey) J. Agardh Lobophora variegata (Lamouroux) Womersley Lobospira bicuspidata Areschoug Pachydictyon paniculatum (J. Agardh) J. Agardh Padina australis Hauck Padina crassa Yamada Padina frasert (Greville) Greville Padina tenuis Bory Spatoglossum macrodontum J. Agardh Stypopodium flabelliforme Weber-van Bosse Taonza australasica J. Agardh Zonaria angustata (Kuetzing) Papenfuss Zonaria crenata J. Agardh Zonaria diesingiana J. Agardh The most abundant species, in terms of cover and biomass, 1n the Sydney area is Zonaria diesingiana. Some species are apparently rare (Padina australis) or rare but nonetheless widely distributed in New South Wales (Zonaria crenata). Other species have more restricted distributions in southern New South Wales (Lobospira bicuspidata, Pachydictyon paniculatum, Zonaria angustata) though they are otherwise more widely dis- tributed in southern Australia. Some collections made in the course of this study were new records, e.g. Homoeostrichus olsenit (reported in Womersley, 1987) and Distromium flabellatum. Three genera are present in southern Australia but absent from New South Wales: Scoresbyella, Chlanidophora and Glossophora. The record of Glossophora nigricans from Plantation Point, New South Wales (May and Larkum, 1981) is of Dilophus intermedius. Only one genus, Stypopodium, is present in New South Wales but absent on the southern Australian coast. Only two genera (Homoeostrichus and Lobospira), occur on the New South Wales mainland coast but not at Lord Howe Island, but all the genera at Lord Howe Island are found on the mainland coast. KEY TO THE DICTYOTALES IN NEW SOUTH WALES ib Growth initiated by aysinglerapicalicellt 3 tt ota ay ene ee 2 1* Growth initiated from a marginal row or cluster of apical cells .............. 4 Ze Thallus with a single cortical layer and a single medullary layer throushoutia iersek atitesd coal aoe hes ate eens Dictyota PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 10% We. ae P. A. FARRANT ANDR. J. KING 373 Thallus with more than one layer of either cortical cells oninedullarvacellissatleast alomgennan cams eis ee) ee eins cpaia yer eens iene 3) Medulla single-layered, cortex two or more cells thick in oldermthallusipantssagleasiatthe marginser eee ee eae ee Fachydictyon Medulla two or more cells thick, at least at the margins, COMI SuANeSenertsal Hao WMONE od sep bias oo noo dnudbeeodaoacoe be Dilophus Mienistempailocalizediclusterottapicalucellsieraser cen. Sea eee 5 Mreristemnofcerminallrowotapicaliccellissyiams seen) se era oar eee 7 Thallus with thickened terete axis and slender flat branches with ultimate divisions spirally twisted and bicuspidate ............. Lobospira sbivallwslattenedithnowe homies eis tae een) ney i ee aes ee See eee 6 ROndawitheaspromilment milli \nencr peer) went) keen eee ec eee Dictyopteris Erondawithoutasprominentamidniloies . 0. whee t.et. seins een eee Spatoglossum AMICAMaArciMnbontaethalluspnnollediasstirecst: “reine e ie Leer ian eee Padina picalomarcimolsthertiallusimotimnollediae wn.) 6-0 eee ee 8 sshvalllusstwoycellsithickethroughoutes 4.12 naa ae ee Distromium Thallus more than two cells thick, at least in mature parts .................. g Medullary cells rectangular and in regular tiers in transverse section ........ 10 Medullary cells neither uniformly rectangular nor in regular KIETS PMA Von ATS VELSE sSeCtlOM gees iateh (seed yale dys peice he ena A, A Ry 12 Central medullary cells distinctly larger than other medullary Cellistinatiral SVETSe Se CLIO muncrsne <7 Ait. te haienetacs Mn reeteine artes ede Me el Lobophora Medullary cells of similar size throughout transverse section ............... 11 Generally paired cortical cells to each medullary cell, sporangia producing cightaplanosponesimerie 4.) sco aah AR ene cra ora: Zonaria Generally single cortical cell to each medullary cell, sporangia PLoducinetouleaplanosponres\epiae we oe een aera: Homoeostrichus Pronounced size differentiation between smaller pigmented cortical cells and larger colourless medullary cells, distal fronds thick, becoming 4 cells thick several cells below the apex .......... Stypopodium No pronounced difference in size between pigmented cortical cells and colourless medullary cells, distal fronds thin, remaining 2 cells thick for some distance below the apex ............... Taonia Dictyota Lamouroux Growth initiated by a single relatively large apical cell. Thalli complanate, membranous; fronds with dichotomous or partly pseudo-alternate branching; hair tufts not associated with reproductive organs; in section a single layer of large medullary cells and a single layered cortex of small pigmented cells. There are three species of Dictyota on the New South Wales coast, D. alternifida, D. bartayresui and D. dichotoma. KEY TO THE SPECIES OF DICT YOTA IN NEW SOUTH WALES M, lis ye 2S Fronds broad at the apices, tapering towards the base ............ D. bartayresi Eronds lincear.o1ntapenne slightly towardsthelapex.) (6) ase ee 2 Fronds narrow, generally about 1 mm wide (range 0.5-4mm [2-4mm wide forms have 2-4cm long internodes and the thallus tends to spiral] )....... D. alternifida Fronds broad, generally greater than 4 mm wide (range 2-14mm [2-4mm forms have internodes shorter than 2cm and the thallus does not tend to Spiral} te user encis SeeeToeeece saver oben) aa ages D. dichotoma PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 374 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Dictyota alternifida J. Agardh, 1894:80. Womersley, 1987: 198. 5cm Fig. 2. Dictyota alternifida: (A) habit of broad-fronded open coast form, slightly spiralled (UNSW 17605a), (B), habit of fine form (UNSW 18050), and (C) usual habit (UNSW 17779). Thalli (5-)7-10(-14)cm long; fronds 0.5-2(-4)mm wide (Fig. 2A-C), linear, some- times twisted, tapering slightly toward the tip, apices usually acute, margins smooth to undulate (occasionally with proliferations); branching dichotomous at 0.5-1.5(-4)cm PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 31D intervals, with a narrow branching angle (less than 90°, usually 45° or less); sporangia scattered (Fig. 2A). Plants from rough-water localities have wider fronds (to 4mm), but are distin- guished from Dictyota dichotoma by their longer internodes (2-4cm) and the tendency of the thallus to spiral (Fig. 2A). Such plants form a continuous series with more typical D. alternifida specimens and can be readily distinguished from D. dichotoma in the field. The wide variation in this species and the relationship to other ‘narrow’ species requires detailed study. Plants referred to D. alternifida from New South Wales coastal saline lagoons display unusual thallus variations including frond proliferations, distal crowd- ing of branches, recurved branching angles, apiculate branch ends and spiralling. One unusual specimen from the Coffs Harbour region (UNSW 18605) is here referred to D. alternifida, although this may prove to be a different species should more specimens become available. Seasonality: collected in all months; sporangial plants in all months except Janu- ary; no sexual material found. Australian distribution: not previously recorded for New South Wales, although Lucas (1909) recorded the species for northern and southern Australia; the most common species of Dictyota in southern Australia (Womersley, 1987); found right along the New South Wales coast in a range of habitats, from shallow sandy sublittoral areas on rough water coasts to positions of extreme shelter in coastal saline lagoons. Selected specimens examined: New South Wales: Hastings Point, 19-v-1986, Phillips, MUCV 2562; Coffs Harbour, 2-4m, 4-x-1981, Farrant, UNSW 18050; Hat Head, LWM, 9-x-1985, P and W. Farrant, UNSW 18643; Smiths Lake, 23-i1v-1984, King, UNSW 16204; Long Reef, 10-vii-1975, Harada R2101, NSW; Fairy Bower, 2-3m, 1-xu- 1985, Farrant, UNSW 18594; Dobroyd, 10m, 24-ix-1985, Farrant, UNSW 18530; Clovelly, 1-3m, 28-11-1985, Farrant, UNSW 17659. South Australia: Cape Northumberland (38°03’S, 140°40’E), 15-x-1985, Womersley, NSW ex ADU A56812. Womersley (1987) has noted that species concepts in the genus Dictyota are not well established and that the degree of variation in many species is considerable. Dizctyota linearis J. Agardh and D. dichotoma var. intricata [both recorded for New South Wales by May (1939) ] are here placed under D. alternifida. Dictyota linearis is characterized by narrow twisted fronds. The majority of the plants from New South Wales do not show this feature, although many fit the description of the species given by Earle (1969) (who nonetheless questions the validity of this species). Ductyota furcellata (C. Agardh) J. Agardh was recorded by May et al. (1978) and by Borowitzka et al. (1982) for New South Wales. All specimens on which these records were based are considered to be D. alterni- fida. We have also placed under D. alternifida all narrow forms, including those from coastal saline lagoons. Dictyota bartayresa Lamouroux, 1809:331. Cribb, 1954:20; Allender and Kraft, 1983:112. Thalli 5-10cm long; fronds 4-5mm wide at the apices, tapering to 2mm or less at the base; branching sub-dichotomous, dichotomies frequently developing unevenly, branching angle usually 70-90° (Fig. 3A-C); sporangia with involucral cells. Dictyota bartayresii plants in New South Wales exhibit broad obtuse apices and narrow bases as noted by Cribb (1954). The species is extremely variable (Cribb, 1954) and Allender and Kraft (1983) placed plants from Lord Howe Island (which taper to the apices as well as the base) into this taxon. The well-developed striations reported by Allender and Kraft (1983) and seen in some Queensland specimens (e.g. UNSW 15354) are not seen in New South Wales plants. PROG. LINN. SOG. N.S.W,, 110 (4), (1988) 1989 376 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Seasonality: may be seasonal in New South Wales since most plants collected in the period September-November; only one sporangial plant examined (NSW: V. May 1045, Collaroy). Australian distribution: pantropical, including south-eastern Queensland (Cribb, 1954) and Lord Howe Island (Allender and Kraft, 1983); now recorded for New South Wales, on rocks, 5-15m deep. Selected specimens examined: New South Wales: Broughton I., 15m, 28-iv-1985, Farrant, UNSW 18017; Collaroy, drift, 16-xi-1945, V. May 1045, NSW; Long Reef, 15m, 16-xi-1985, Farrant, UNSW 18575; Fairlight, 2-3m, 16-1x-1985, Farrant, UNSW 18511; Dobroyd, 10m, 24-ix-1985, Farrant, UNSW 18531; Clovelly, 27-vi-1987, D. May, MUCV 2576; Bare I., 10m, 17-x-1985, Farrant, UNSW 18564; Plantation Point, 1-x- 1974, Larkum and May, NSW. Dictyota dichotoma (Hudson) Lamouroux, 1809:331. Womersley, 1987:194. Thalli 3-20cm long, often with a blue-green irridescence im situ; fronds 5-10 (-14)mm wide, linear, tapering slightly towards broad apices, without proliferations; branching dichotomous, branching angle 15-45° (Fig. 4A-D); sporangia (Fig. 4E) scat- tered, often densely, over both frond surfaces but frequently in broad transverse bands (Fig. 4A), best seen in plants 7m situ; sexual plants dioecious, oogonia (Fig. 4B,F) and antheridia (Fig. 4C,G) in scattered sori; male plants often recognizable from the more or less circular ‘patches’ surrounded by involucral cells which remain after the antheridia have been released. Seasonality: fertile plants year-round; sporangial plants in every month except May; gametangial plants in all but May, August and November (King and Farrant, 1987). Australian distribution: widely distributed (Womersley, 1967), with numerous records for New South Wales (Gepp and Gepp, 1906; Lucas, 1909, 1914; May, 1939; Cribb, 1954; May et al., 1970); present in bays, estuaries and on rough water coasts at depths to 16m, on rock or other substrata, often epiphytic. Selected specimens examined: New South Wales: NW Solitary I., 10-16m, 6-x- 1985, Farrant, UNSW 18613; Coffs Harbour, 28-viii-1980, Millar, MELU AM 362; Port Macquarie, 0-2m, 10-x-1985, P and W. Farrant, UNSW 18658; Newport, 21-x-1944, V. May 422, NSW; Clovelly, 2-3m, 10-iv-1985, Farrant and Puttock, UNSW 17760; Botany Bay, Jan. 1905, Lucas, NSW 140397; Bare I., 1-2m, 13-11-1985, Farrant and Puttock, UNSW 17692; Kiama, 20-xi-1945, V. May 1062, NSW; Crookhaven Heads, 30-iv-1977, King and Kertesz, UNSW 15347. The New South Wales records of Dictyota papenfussi: Lindauer (Lindauer et al., 1961) and of D. radicans Harvey (May, 1939) are here referred to D. dichotoma. Pachydictyon J. Agardh Growth initiated by a single protruding apical cell. Thalli dichotomous, develop- ing axes which bear alternate lateral branch systems; branches compressed (Womersley, 1987); in section consisting of a single-celled medulla, and a multilayered cortex in older, lower axes. Pachydictyon paniculatum (J. Agardh) J. Agardh 1894:84. Womersley 1987:211. Thalli 6-10cm long, attached by numerous flattened slender outgrowths from near the base; fronds 0.5-lmm wide in upper parts, lower parts 2-3mm wide; upper branches fastigiate, subdichotomous to lateral (Fig. 5A,B); hair tufts and sporangia scattered. PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING ST ~ & baae 5cm \ 56 GH CUTHVIOIT OU UOSVE WOO OHGoOBsCe 100um AL Fig. 3 (above). Dictyota bartayresi: (A) habit of large plant (UNSW 18554), (B) and (C) habit of small plants (UNSW 18511). Fig. 4 (below). Dictyota dichotoma: (A) habit of sporangial plant (UNSW 18259), (B) habit of female plant (UNSW 18069), (C) habit of male plant (UNSW 17748), (D) small plant (UNSW 17677), (E) TS. sporangial plant (UNSW 18259), (F) T.S. female plant (UNSW 18069), and (G) TS. male plant (UNSW 18038, wet material only). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 378 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES 5cm Fig. 5. Pachydictyon paniculatum: (A) habit (V. May 2040, NSW), (B) frond apex. New South Wales specimens are smaller than those described from more extensive collections in southern Australia. Seasonality: early collections made in January (1943, 1946); however no specimens collected in this study. Australian distribution: in New South Wales appears to be confined to the south coast where it is rare; earlier reports of the species extending as far north as Sydney (e.g. Borowitzka et al., 1982) probably based upon two specimens of Dictyota in NSW collected by Lucas and incorrectly identified; in southern Australia epiphytic on larger algae in the upper subtidal under moderate to strong water movement (Womersley, 1987). Selected specimens examined: New South Wales: Moruya, 7-1-1943, V. May 985, NSW; Ulladulla, 13-1-1946, V May 2040, NSW; Green Cape, 1970, Ducker and King, MELU 20694. Victoria: Flinders, 5-1-1976, King, UNSW 16049. COMPARISON WITH PACHYDICT YON SPECIES IN ADJACENT REGIONS A second species, Pachydictyon polycladum (Kuetzing) Womersley, occurs from Champion Bay, Western Australia to Port Phillip Bay, Victoria (Womersley, 1987), but has not been recorded in New South Wales. A third species of Pachydictyon has been recorded for Lord Howe Island. This species, P aegerrime Allender and Kraft, just qualifying as a Pachydictyon because on its multicellular cortex, was described on the basis of a single plant, indistinguishable in the field from Ductyota bartayresiu but lacking horizontal banding, spiralling and involucre cells (Allender and Kraft, 1983). Dilophus J. Agardh Growth from a single apical cell. Thalli erect, complanate, dichotomously branched or with irregular lateral branching; fronds in section exhibiting a single-celled cortex and a single-celled medulla which becomes multilayered at least at the margins. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 379 FEL Ext Sl? = \, s 4 Rte m is izeeeze J Vite ( J Tahar : 200um {| || 100um 7 anen COodoCOOANdodHABONGeOGODNHOGGE Fig. 6. Dilophus intermedius: (A) habit of sporangial plant (UNSW 13789), (B) T.S. frond margin (UNSW 18602), (C) sporangia (UNSW 17708), (D) male gametangia (UNSW 18338). Two species of Dilophus are common on the New South Wales coast: D. intermedius and D. marginatus. KEY TO THE SPECIES OF DILOPHUS IN NEW SOUTH WALES I, Mature fronds with branches more than 6mm (usually about 10mm) wide, sori in block-like patches, branches often with abundant flattened proliferations, branchespnoticonnugated meer tenes ret eu atHy seaiin eane D. intermedius 1*. Mature fronds with branches less than 6mm wide, sori scattered, not in block-like patches, branches without proliferations, branches corrugated .... D. marginatus Dilophus intermedius (Zanardini) Allender and Kraft, 1983:118. Thalli 10-40cm long, attached by wiry stolons (Fig. 6A); branches 8-13mm (usually about 10mm) wide, often bearing proliferations up to 10mm long in rows across lower PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 380 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES frond surfaces; in transverse section medulla 1(-2) cells thick, up to 4-6 cells thick at the margins (Fig. 6B); sporangia (Fig. 6C) in block-like patterns on the frond surface (Fig. 6A); antheridia in indusiate sori (Fig. 6D). The species is apparently very variable (Allender and Kraft, 1983: fig. 25) and we include the collection from Long Reef (MELU GK6921, duplicate in UNSW) within this species. Plants in this collection are dark brown, with diffuse sori rather than block- like sori, and proliferations along the margins which may be consequent on damage. Seasonality: sporangial plants collected in March, April, May, June and October; male plants found in March and April, the first record of gametangial plants of Dilophus intermedius. Australian distribution: Lord Howe Island, and the mainland coast from Sydney north to Queensland (Allender and Kraft, 1983; Cribb, 1954; Lindauer et a/. 1961; May, 1939; Gepp and Gepp, 1906); in this survey as far south as Jervis Bay, growing on rocks in the upper sublittoral. Selected specimens examined: New South Wales: Hastings Point, 19-v-1986, Phillips, MUCV 2558; Split Solitary I., 16m, 5-x-1985, Farrant, UNSW 18602; Coffs Harbour, 18-1v-1980, Millar and Mix, MELU AM 061; Seal Rocks, 0-2m, 10-x-1985, P and W. Farrant, UNSW 18683; Tuggerah Lakes, Apr. 1911, Lucas, NSW A1351; Long Reef, 12-vi1-1979, Kraft and Borowitzka, MELU GK 6921 (duplicate in UNSW: 18400); Fairlight, 1-2m, 20-11-1986, Farrant, UNSW 18338; Fairlight, 1-3m, 14-v-1985, Farrant, UNSW 18055; Long Bay, July 1903, Lucas, NSW 140425; Jervis Bay, drift, 18-11-1985, King, UNSW 17708. Queensland: Caloundra, Jan. 1909, Lucas, NSW 140451. Tir tit Vc ax poceay Cs alaladnts,',_\) Bie area) 100um | COOGI00 100um Cc Fig. 7. Dilophus marginatus: (A) habit of sporangial plant, with sporangia concealing the wrinkled nature of the thallus (UNSW 18225), (B) T.S. frond margin (UNSW 18042), (C) sporangia (UNSW 18355). Dilophus marginatus J. Agardh, 1894:91. Allender and Kraft, 1983:118; Womersley, 1987:202. Thalli 5-18cm (usually about 10cm) long, basally stoloniferous (Fig. 7A); fronds 3- PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 381 6mm wide, linear, mature blades corrugated due to transverse wrinkling; hair tufts scattered; in transverse section a single-celled medulla becoming 2(-3) cells thick at the frond margins (Fig. 7B); sporangia (Fig. 7C) scattered on the frond surface, especially abundant in the corrugations. Seasonality: present in all months, although abundance was seasonal, peaking in autumn-winter (King and Farrant, 1987); sporangial plants found in all months except December and January; no gametophytic plants known for this species. Australian distribution: from South Australia to Queensland, and from Lord Howe Island (May, 1939; Cribb, 1954; Allender and Kraft, 1983; Womersley, 1987); common in shallow sublittoral rocky and sandy habitats and lower eulittoral pools. Selected specimens examined: New South Wales: Hastings Point, 19-v-1986, Phillips, MUCV 2557; NW Solitary I., 10-16m, 6-x-1985, Farrant, UNSW 18606; Port Macquarie (31°27’S, 151°26’E), 21-xi-1972, Coveny 4737, NSW; Broughton I., 15m, 28- iv-1985, Farrant, UNSW 18016; Fairlight, 1-2m, 20-v-1986, Farrant, UNSW 18372; Clovelly, 1-3m, 28-11-1985, Farrant, UNSW 17656; Long Bay, June 1915, Lucas, NSW; Steamers Beach, 11-vii-1979, King and Kertesz, UNSW 15363. Victoria: Brighton, Jan. 1900, Lucas, NSW. COMPARISON WITH DILOPHUS SPECIES IN ADJACENT REGIONS Species concepts in the genus Dilophus are unsatisfactory. Womersley (1987) has reduced the number of species on the southern Australian coast from eleven (Womers- ley, 1967) to seven: D. marginatus, D. robustus (J. Agardh) Womersley, D. angustus J. Agardh, D. tener J. Agardh, D. crinitus J. Agardh, D. fastigiatus (Sonder) J. Agardh, and D. gunnianus J. Agardh. ‘Two of the species of Dzlophus recorded for northern Australia, D. intermedius and D. marginatus (Lewis, 1985), are found in southern Queensland and at Lord Howe Island (Allender and Kraft, 1983). Lobospira Areschoug Growth apical from short rows of cells at the apices of the axes and laterals (not in the determinate laterals). Thalli with recurved attaching lower lateral branches, upper free branches twisted and bearing bicuspid determinate laterals; midrib present, especially in the lower half of branches; sporangia scattered and sunken in the thallus. The monotypic genus Lobospira is easily distinguished from all other genera by its distinctive morphology. Growth 1s from a cluster of apical cells (Edelstein and Womers- llevan Oi75)): Lobospira bicuspidata Areschoug, 1854:364. Edelstein and Womersley, 1975:149; Womersley, 1987:214. Thalli 10-25cm long; fronds 1-2mm wide (Fig. 8A,B). Seasonality: only one collection from New South Wales, that of Lucas in January 1910. Australian distribution: southern Australia (from Nickol Bay, Western Australia to Eden, New South Wales) (Womersley, 1987); found from just below LWM to a depth of 13m; only collected once in New South Wales (Lucas, Eden, Jan. 1910, 10 specimens in NSW) so possibly drift from the south; not collected in this survey. Selected specimens examined: New South Wales: Eden, Jan. 1910, Lucas, NSW. Victoria: Point Lonsdale, 26-11-1979, King, UNSW 16018; Sandringham, Jan. 1900, Lucas, NSW. South Australia: Port MacDonnell, 13-iv-1975, Harada R2666, NSW. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 382 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Nee she 5cm Fig. 8. Lobospira bicuspidata: (A) habit (Eden, Jan. 1910, A. H. S. Lucas), (B) frond apex. Dictyopteris Lamouroux Growth from a small group of initials generally in a slight apical depression. Thalli erect, smooth, flattened, membranous to coarse, mostly delicate; fronds with a conspicuous midrib, branches arising from the midrib or margins; hair clusters and reproductive organs on both surfaces; in section consisting of an inner zone of more or less cuboidal to angular colourless cells, and an outer layer of small cuboidal pigmented cells. Dictyopteris arostichoides is the only species of Dictyopteris on the New South Wales coast. Dictyopteris acrostichoides (J. Agardh) Boergesen, 1935:36. Womersley 1987:226. Faliserts acrostichoides J. Agardh, 1882: 133. Thalli 15-30(-55)cm long, branching often but not always from or near the midrib, with some marginal branching in upper parts; fronds (0.4-)0.8-1.0(-1.5)cm wide (Fig. 9A); reflexed lines of hair tufts present, no veinlets; 4(-6) cells thick except in the region of the midrib (Fig. 9B-D); sporangia embedded in the cortical layer (Fig. 9B,C), scattered in two distinct elongate sori, leaving the midrib area and marginal zones sterile (Fig 9A); fertile regions of male plants forming irregularly shaped pale patches on PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 383 4 od oS DY Va DORI OE Fig. 9. Dictyopteris acrostichoides: (A) habit of sporangial plant, showing longitudinal sori (arrow) (UNSW 17663), (B) T.S. of frond showing hair tufts (UNSW 18345), (C) T.S. of frond showing sporangia (UNSW 17647), (D) T.S. of male frond showing midrib and antheridial sori (UNSW 17647). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 384 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES the frond wings with antheridial sori sunken in the thallus (Fig. 9D); in dried herbarium material male plants difficult to distinguish from sterile plants. Seasonality: collected in all months of the year; sporangial plants in all months except May and August; male gametophytes (Fig. 9D), previously unrecorded, collected in February and September from Sydney Harbour; female plants not found. Australian distribution: from Warrnambool, Victoria, to Rockingham, Queens- land, and along the northern coast of Tasmania (Womersley, 1987); many published records for New South Wales (e.g. Lucas, 1913, 1914; May, 1939; Womersley, 1949), where plants grow in the shallow sublittoral, in sheltered localities on rough water coasts and in bays. Selected specimens examined: New South Wales: Pottsville Beach, 19-v-1986, Phillips, MUCV 2560; Woolgoolga, Jan. 1972, V May, NSW 126974; Crescent Head, LWM, 9-x-1985, P and W. Farrant, UNSW 18652; Camp Cove, 1-2m, 28-11-1985, Farrant and Puttock, UNSW 17647; Lady Jane Beach, 1-3m, 28-11-1985, Farrant and Puttock, UNSW 17663; Mrs Macquarie’s Point, 1-2m, 13-1x-1985, Farrant, UNSW 18291; Point Piper, July 1899, Lucas, NSW; Bare I., 6m, 25-vii-1985, Farrant, UNSW 18246; Crook- haven Heads, LWM, 12-iv-1986, King, UNSW 18635. Queensland: Noosa, Nov. 1943, V. May 989, NSW; Margate, 21-x11-1943, Vv May 987, NSW; Sandgate, Dec. 1913, Lucas, NSW. COMPARISON WITH DICTYOPTERIS SPECIES IN ADJACENT REGIONS Four species of Dictyopteris (D. australis (Sonder) Askenasy, D. gracilis Womersley, D. muellert (Sonder) Reinbold, and D. nzgricans Womersley) occur in southern Australia but not New South Wales (Womersley, 1987). There are five species recorded for Lord Howe Island: D. australis and D. plagiogramma (Montagne) Vickers, both of which have veinlets; D. repens (Okamura) Boergesen and D. delicatula Lamouroux, distinguished by their small size (and perhaps representing a single species); and D. crassinervia (Zanardin1) Schmidt distinguished by its unique winged apices (Allender and Kraft, 1983). Dictyop- teris crassinervia, D. delicatula and D. repens are also recorded from Queensland (Allender and Kraft, 1983; Ngan and Price, 1979, 1980). REJECTED SPECIES RECORDS Dictyopteris australis (Sonder) Askenasy: Allender and Kraft (1983) stated that D. australis ‘is now known to occur generally around Australia? There are in fact no records for Victoria or New South Wales though the species occurs in South Australia, Western Australia, Queensland (Womersley, 1987) and at Lord Howe Island (Allender and Kraft, 1983). The species is similar to D. acrostichoides, but is easily distinguished by the presence of fine lateral veins and by the distribution of the sori and associated hair tufts in reflexed lines. The frond wings of D. australis are generally distromatic rather than of four or more cells as in D. acrostichordes. Dictyopteris muellert (Sonder) Reinbold: Harvey’s specimen of D. mueller: (NSW 140374, Harv. Alg. Austr. Exsic. n. 87, 1855) was referred to by May (1939) and on that basis Womersley (1987) recorded D. muellert for New South Wales. The specimen has no collecting locality. Harvey (1860) listed Port Jackson, New South Wales as a collecting locality but he stated that the species was rare on the east coast. The species record for New South Wales therefore has no reliable herbarium voucher specimen and the species has not been recorded otherwise from New South Wales. Dictyopteris muellert has a southern Australian distribution and is easily distinguished from Dictyopteris acrostichoides by the evenly scattered hair tufts which are not arranged in reflexed lines; fertile D. mueller: material is easily recognized, since sporangia, oogonia and antheridial sori PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P. A. FFRRANT ANDR. J. KING 385 occur in broad bands across the thallus and the midrib and marginal regions are sterile (J. Phillips, pers. comm. ). Dictyopteris plagiogramma (Montagne) Vickers: although Allender and Kraft (1983) recorded this species from the Coffs Harbour region of New South Wales, there is no herbarium voucher specimen (A. Miller and G. Kraft, pers. comm.). Dictyopteris plagiogramma occurs at Lord Howe Island (Allender and Kraft, 1983) and in northern Australia (Lewis, 1985). It is distinguished from D. acrostichoides by the presence of fine lateral veins and by the arrangement of hair tufts in single longitudinal rows on each side of the midrib. Dictyopteris woodwardu (R. Brown ex Turner) Schmidt: was recorded for Ballina by Sonder (1871) as Halyseris polypoidioides var. woodwardia. Vhis was based on a specimen in MEL (584114) which has been annotated by B. M. Allender (12-vu-1981) as D, acrostichoides. Dictyopteris woodward is otherwise recorded for northern Australia (Lewis, 1985). The species is distinguished from D. acrostichoides by having frond margins fringed with minute spine-like teeth. Spatoglossum Kuetzing Growth initiated by a group of apical cells. Thalli broad, complanate; fronds branching irregularly, hair tufts scattered; sporangia scattered and sunken in the frond surface. Spatoglossum macrodontum is the only species of the genus in New South Wales. CORA RErAnooaoor eee ie a eee WOOO oa Ceo 100yum Fig. 10. Spatoglossum macrodontum: (A) habit of small plant (UNSW 17740), (B) habit of large broad plant (UNSW 18714), (C) TS. frond (UNSW 18251), (D), section through sporangia (UNSW 17791). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 386 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Spatoglossum macrodontum J. Agardh, 1882:113. Allender and Kraft, 1983:100. Thalli 10-35cm long, medium to dark brown (drying to a greenish-brown colour); fronds 3-8cm wide, complanate; branching irregular with unequal development of dichotomies, margins smooth in young plants (Fig. 10A) but irregular to dentate in older, larger plants (Fig. 10B); 4-6 cells thick and cells arranged irregularly (Fig. 10C); sporangia scattered and sunken in the frond surface (Fig. 10D). Seasonality: collected in all months except February, but seasonally abundant, with young plants present at the end of winter and early spring; sporangial plants collected in most months in this study and spent male gametangial plants in June, July and August. Australian distribution: Jervis Bay, New South Wales, north to Queensland and the Northern Territory, and at Lord Howe Island (Allender and Kraft, 1983; Lucas, 1909, 1914; May, 1939); locally abundant in the shallow sublittoral. Selected specimens examined: New South Wales: Port Stephens, June 1909, Lucas, NSW 140350; Port Stephens, 0-2m, 11-x-1985, P and W. Farrant, UNSW 18714; Vaucluse, 1m, 21-x1-1985, Farrant, UNSW 18577; Mrs Macquarie’s Point, 2-3m, 29-vii- 1985, Farrant, UNSW 18251; Mrs Macquarie’s Point, 2m, 27-vi-1985, Farrant and Puttock, UNSW 18208; Jervis Bay, 9-x-1941, V. May 983, NSW. Queensland: Magnetic I., 15-v- 1987, Phillips, MUCV 2565; Redcliffe, 5-vi-1980, Alderslade R3988, NSW. COMPARISON WITH SPATOGLOSSUM SPECIES IN ADJACENT REGIONS Spatoglossum macrodontum as well as two other species, S. asperum J. Agardh and S. shroederr (Mertens) J. Agardh, have been recorded from northern Australia (Lewis, 1985). Spatoglossum macrodontum is the only species found at Lord Howe Island (Allender and Kraft, 1983). Spatoglossum macrodontum appears to be closely related to S. australasicum Kuetzing from South Australia, but there is little material (and no fertile material) of S. australasicum available for comparison (Womersley, 1987). Padina Adanson Growth from a marginal row of apical cells. Thalli flabellate with inrolled margins, calcified to various degrees. Four species of Padina occur on the New South Wales coast: Padina australis which 1s not common, P. crassa, P. fraser, and P. tenuis. The four species, especially the last two, are difficult to separate without sectioning and/or reproductive material (Fig. 11). KEY TO THE SPECIES OF PADINA IN NEW SOUTH WALES 1. Thallus),2 cells thick«throughout.:.4 2b ear ase ete, ite ec eee Bee 2 Le vhallusimore than 2 cellsithick except near the|apex (4. sacar oe 3 2 Sporangia on upper surface in concentric lines bordered on each side by hair bands that alternate on upper and lower surfaces; sporangial regions separated from each other DypSienile sz OleSe awe eu cus id Bun ae cater Oe eae P. australis 2.* Sporangia on upper surface in concentric lines bordered on each side by hair bands that are only on the upper surface; sporangial regions not separated from each other by Tied eae ZO) ga) ee ee OS an, eR Re Re Ve ae Mee ON EE or P. tenuis D: halls /Jicelisainickathnoughovitern tyasieiocycaacua akan ce P. frasert 2 Ne WGhrallisiG=1Opcellstthitc Ke jesitkecai tht ke Eee eee ae ee ae P. crassa PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 387 upper = inner P. australis P. crassa P. fraseri P. tenuis Fig. 11 (above). Diagram showing distribution of hairs and sori on New South Wales species of Padina (not to scale). Fig. 12 (below). Padina australis: (A) habit of gametangial plant (UNSW 17705), (B) TS. top of frond (UNSW 17705), (C) section through female and male gametangia (UNSW 17705). PROC. LINN. SOG. N.S.W., 110 (4), (1988) 1989 388 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Padina australis Hauck, 1887:44. Allender and Kraft, 1983:85. Thalli 6-10cm long; fronds 4-12cm wide (Fig. 12A), with conspicuous hairs on both surfaces, hair lines 1-1.5mm apart; distromatic throughout (Fig. 12B); gametangial plants monoecious, with female and male gametangia in mixed sori (Fig. 12C) on the upper (inner) frond surface, alternating fertile and sterile zones. Seasonality: collected in February and March only, from Church Point, Pittwater; gametangial plants present in both collections; no sporangial plants found. Australian distribution: reported from Lord Howe Island, and in mainland Aus- tralia from the central New South Wales coast to the Northern Territory (Allender and Kraft, 1983); in this survey found on rocks just below low water mark in a very sheltered part of the estuary. Selected specimens examined: New South Wales: Lord Howe I., 14-v-1977, Kraft, MELU A037276; Lord Howe I., 16-xi1-1986, Kraft and Millar, MELU A037275; Church Point Pittwater, LWM, 17-11-1985, Farrant, UNSW 17705. Padina crassa Yamada, 1931:67. Allender and Kraft, 1983:87. Thalli 6-16cm long; fronds 3-10cm wide (Fig. 13A), hairs conspicuous, in alternat- ing rows on each frond surface, (2-)4-5(-7.5)mm apart; fronds 6-8(-10) cells thick except at the apex (Fig. 13B,C); sporangial sori indusiate (Fig. 13B) and mainly on the upper (inner) side of the frond (occasionally on both surfaces), fertile zones not separated by sterile zones. Padina crassa specimens from the New South Wales mainland coast agree with Allender and Kraft’s (1983) Lord Howe Island material in that they have no sterile zones, have the same seasonality and indusiate sori. The mainland material has hairs on both frond surfaces whereas the Lord Howe Island plants have them only on the upper surfaces. Allender and Kraft (1983) stated that ‘there is little if any significant difference between . . . PR. gymnospora and FP. crassa from the eastern and southern Pacific. Womers- ley (1987) recorded PR. gymnospora (Kuetzing) Sonder in southern Australia, noting that it has a distribution extending into warm temperate waters on both eastern and western coasts of Australia. He noted however that, according to Allender and Kraft (1983), P crassa from Lord Howe Island has sori on the outer (lower) surface only, whereas in P gymnospora the sori are mainly (but not only) on the upper surface. Allender and Kraft (1983) attempted to clarify the confusion which ‘surrounds the terminology used to describe the morphology and anatomy of the species’ of Padina and correctly defined the use of the terms inner and outer, the inner being the ‘side towards which the marginal curl is directed’. In their descriptions of P. australis, P. crassa and P. tenuis Allender and Kraft stated that the sporangial sori are located on the outer frond surfaces, and in their fig. 6 the outer surface is correctly referred to as the ‘upper’ sur- face. We have examined fertile specimens of all these species from Lord Howe Island (MELU A037276, MELU A037275, MELU GK9560, MELU A037274) and of P fraser: from Victoria (MELU 4290), and the sori in all cases occur on the surface to which the marginal roll is directed, 1.e., the inner surfaces. The descriptions of these species in Allender and Kraft (1983) should therefore read sporangial sori on inner frond surface rather than outer. The inner surface is then equivalent to the upper surface in the terminology used by Womersley (1987). The material from New South Wales and Lord Howe Island thus differs from Fadina gymnospora in that it has indusiate sori whereas P. gymnospora has essentially non- indusiate sori. Padina gymnospora and the New South Wales specimens have hairs on both surfaces whereas the Lord Howe Island species has hairs on the upper surface only. The PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P.A. FARRANT ANDR. J. KING 389 Rice eh re oo = Set SSeS ie ie soon | EO ae ag B 100um woes 5cm SEURUASUCGUREES e META) DUCTUS, oop Fig. 13 (above). Padina crassa: (A) habit (UNSW 18041), (B) sporangial sorus (UNSW 17766), (C) base of frond (UNSW 18803). Fig. 14 (below). Padina fraseri: (A) habit (UNSW 18685), (B) L.S. apex (UNSW 18002), (C) T.S. base of frond (UNSW 18685), (D) section through male sorus (UNSW 17665). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 390 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES relationship between these two taxa deserves attention; examination of type specimens may show that the two are not distinct. Seasonality: absent in the winter months; sporangial plants collected in the warmer months (King and Farrant, 1987); no gametangial plants collected. Australian distribution: Lord Howe Island (Allender and Kraft, 1983) and in Queensland (UNSW 16036); in this survey north to Hastings Point and south to Jervis Bay; found in the shallow upper subtidal, on the open coast and in bays and estuaries. Selected specimens examined: New South Wales: Hastings Point, 19-v-1986, Phillips, MUCV 2563; Coffs Harbour, 2-4m, 4-x-1981, Farrant, UNSW 18041; South West Rocks, LWM, 9-x-1985, P and W. Farrant, UNSW 18637; Lord Howe I., 9-xii-1978, Kraft and Ricker, MELU GK 9560; Fairlight, Im, 29-1-1986, Farrant, UNSW 18829; Vaucluse, 1m, 18-1v-1985, Farrant and Puttock, UNSW 17790; Jervis Bay, LWM, 18-iii- 1985, King, UNSW 17709. Queensland: Noosa, 4-x11-1976, King and Kertesz, UNSW 16036. Padina fraseri (Greville) Greville, 1830, synop.:xliv. Womersley, 1987:217. Thalli 6-9.5cm long, often distinctly crinkled; fronds 3-7.5cm wide (Fig. 14A), with alternate concentric hair lines on both sides of the fronds, (2-)4(-7)mm apart; 3 cells thick almost to the apex (Fig. 14B), with central (medullary) cells larger than cortical cells (Fig. 14B-D); sporangial sori indusiate, no sterile zones between hair lines; male gametangial plants with indusiate antheridial sori (Fig. 14D). Fadina pavonica (Linnaeus) Lamouroux has been recorded for Port Jackson, New South Wales (May, 1939) and generally for New South Wales and Lord Howe Island (Lucas, 1909, 1914, 1935). This species differs from P fraseri in not having the central (medullary) cells longer than the outer cells (Allender and Kraft, 1983: fig. 6D,F). The records of PR. pavonica for New South Wales probably mostly refer to P._frasert, since the NSW specimens that have been examined in the present study, and which are 3 cells thick, have longer central cells and a crinkled thallus, and therefore appear to belong to P. frasert. Seasonality: sporophytic plants collected especially in the warmer months, and male plants, the first sexual plants to be recorded for the species, collected in February. Australian distribution: recorded from south-eastern Australia from Warrnam- bool, Victoria, to the mid north New South Wales coast (Womersley, 1987); grows at and just below low water mark, on rocky open coasts. Selected specimens examined: New South Wales: Crescent Head, LWM, 9-x- 1985, PR and W. Farrant, UNSW 18649; Seal Rocks, 0-2m, 10-x-1985; P and W. Farrant, UNSW 18680; Lady Jane Beach, 1-3m, 28-11-1985, Farrant and Puttock, UNSW 17665; Wollongong, Jan. 1912, Lucas, NSW; Kiama, Dec. 1899, Lucas, NSW; Crookhaven Heads, LWM, 20-iv-1985, Farrant, UNSW 18002; Merimbula, 19-vii-1987, D. May MUCV 2575. Victoria: Barwon Heads, 7-11-1969, Ducker, MELU 4290; Apollo Bay (38°46'S, 143°44’E), 23-1-1967, Womersley, NSW ex ADU A31758. Padina tenuis Bory, 1827:590. Womersley and Bailey, 1970: 292; Allender and Kraft, 1983:83. Thalli 6-18cm long; fronds 3-12cm wide (Fig. 15A), without conspicuous hairs below the first hair line, hair scar lines (1.5-)3-4(-6)mm apart; fronds distromatic throughout (Fig. 15B,C); sporangial sori indusiate (Fig. 15B) or have the remains of an indusium, fertile zones not separated by sterile zones. The New South Wales specimens of Padina tenuis agree well with Allender and Kraft’s (1983) description of the species for Lord Howe Island and the southern Great PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P. A. FARRANT AND R. J. KING 391 Ae oN area EDD cerarn iin ouannanNeNONOH Gg HONE B 100um 2 oo TON : ben MAUGHAM TE GUHA TMEAUINEHE C Fig. 15. Padina tenuis: (A) habit (UNSW 18068), (B) T\S. base of frond through sporangial sorus (UNSW 18583), (C) TS. base of frond (UNSW 18583). Barrier Reef, except that their material has non-indusiate sori and the fronds are 2-3 cells at the base. Almost all New South Wales specimens have indusiate sori and are 2 cells thick throughout. Lord Howe Island material examined during the present study (MELU A037274) showed that the remains of indusia were present. The New South Wales and Lord Howe Island specimens therefore appear to belong to the same taxon, P tenuis, although examination of type material would be required to confirm this. Seasonality: collected in all months, sporangial plants in all months except July, August and October; no gametophytic plants present. Australian distribution: recorded for Lord Howe Island and the southern Great Barrier Reef (Allender and Kraft, 1983) but not reported previously for the mainland coast of New South Wales; grows from a depth of several metres to 16m, on both sheltered and open coast shores. Selected specimens examined: New South Wales: Pottsville Beach, 19-v-1986, Phillips, MUCV 2559; NW Solitary I., 10-16m, 6-x-1985, Farrant, UNSW 18608; Lord Howe I., i-xii-1978, Kraft and Ricker, MELU A037274 (=GK 9405); Broughton I., 15m, 28-iv-1985, Farrant, UNSW 18015; Barrenjoey, 15m, 23-i1-1985, Farrant, UNSW 17728; Mrs Macquarie’s Point, 1-2m, 26-11-1986, Farrant and Puttock, UNSW 18341; Long Bay, Apr. 1900, Lucas, NSW. COMPARISON WITH PADINA SPECIES IN ADJACENT REGIONS Padina australis, P. crassa and P. tenuis occur at Lord Howe Island (Allender and Kraft, 1983). Padina boergesenit Allender and Kraft was described on the basis of Lord Howe Island material (Allender and Kraft, 1983). It is rare and has not been recorded for the New South Wales mainland coast, although Allender and Kraft (1983) suggest that it probably occurs in Queensland. Padina tetrastromatica Hauck, which occurs in Queensland, is similar to P._fraseri except that it becomes 4-layered in older thallus parts (Allender and Kraft, 1983: fig. 6E). Padzna frasert is the only species of Padina for most of the southern Australian coast, with three other species now recognized from the western end of that region: P. elegans Koh ex Womersley, P. gymnospora (Kuetzing) Sonder, and P sanctae-crucis Boergesen (Womersley, 1987). Distromium Levring Growth from a marginal row of apical cells. Thalli two cells thick throughout (3 cells thick at hairs). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 392 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES There is only one species of Distromium (D. flabellatum) in New South Wales. The plants are relatively small compared with those reported from southern Australia, and no fertile plants have been seen in the New South Wales material. Therefore the identifi- cation to specific level is provisional since fertile material is required to establish its re- lationship with D. skottsbergi: Levring (Womersley, 1967; Lindauer et al., 1961). 2cm cS s CY OD Gme-eerieeieeae ote OO 5 @ 3 a B 100um Fig. 16. Distromium flabellatum: (A) habit (UNSW 18849), (B) section of frond (UNSW 18873). Distromium flabellatum Womersley, 1967:218. Womersley, 1987:230. Thalli 2-3cm long, slightly blue-green when seen zn sztu; frond(s) 2-3cm wide (Fig. 16A), flabellate; hairs borne singly (Fig. 16B). Seasonality: collected in this study in April and September; no reproductive plants found. Australian distribution: recorded from 7 Mile Beach, north of Dongara, Western Australia and along the southern Australian coast to Deal I. in Bass Strait (Womersley, 1987); now tentatively recorded for New South Wales; in this study collected at only one locality (Fairlight, Sydney Harbour) on the vertical face of a rocky reef at 4-5m depth; may be more widespread, but is easily overlooked because of its superficial similarity to young Zonaria diesingiana plants, which however differ by being 6-8 cells thick. Selected specimens examined: New South Wales: Fairlight, 5m, 29-iv-1986, Farrant, UNSW 18849; Fairlight, 4m, 25-1x-1986, Farrant, UNSW 18873 (wet material only); South Australia: Aldinga (35°20’S, 138°28’E), 17-ix-1966, Womersley, NSW ex ADU A30711. COMPARISON WITH DISTROMIUM SPECIES IN ADJACENT REGIONS Distromium flabellatum is distinguished from the Lord Howe Island species, D. didymothrix Allender and Kraft, by having hairs borne singly, rather than in pairs derived from a single cell (Allender and Kraft, 1983). There are two species of Distromium in PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 393 southern Australia: D. flabellatum and D. multifidum Womersley. The latter has linear, subdichotomous branches with small sporangial sori in the upper concave parts; the thallus lacks prominent zones of hairs which the larger southern Australian D. flabellatum plants have (Womersley, 1987). Lobophora J. Agardh Growth from a marginal row of apical cells. Thalli flabellate or irregularly lobed, erect or prostrate; fronds in section having medullary cells in regular tiers, with the central layer of cells larger. One species, Lobophora variegata, has been recorded for New South Wales. pend pacowssaaaad s IBAOLE |) os) (a (=) om) cS) sae ee Sale 2 2 eae ae Oe eee) PROCNSSRRSLOLT PA ee OOS IORI OSU CSI IATS SGU FG GEICO SG OOOSGGTOIGHG g a ee a a aU a a a @ ad Fig. 17. Lobophora variegata: (A) habit of broad erect form (UNSW 18018), (B) habit of narrow form (UNSW 17637), (C) TS. of sporangial frond (UNSW 18029), (D) L.S. frond (UNSW 18018). Lobophora variegata (Lamouroux) Womersley, 1967:221. Allender and Kraft, 1983:81. Gymnosorus variegatus (Lamouroux) J. Agardh 1894:11; Pocockiella varvegata (Lamouroux) Papenfuss 1943:467 Thalli 3-11cm long; fronds 3-6cm wide (Fig. 17A,B); in transverse section the medulla usually 9 cells thick (Fig. 17C), less near the apex (Fig. 17D), cells arranged in PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 394 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES regular tiers with the central cells larger than the outer cells; sporangia in isolated sori on the fronds. Young Lobophora variegata plants may be confused with young Zonaria diesingiana plants, but the latter lack the central layer of larger medullary cells. Seasonality: sporangial plants collected in all months; gametangial plants not present in collections. Australian distribution: tropical and temperate coasts of Australia (Womersley, 1987), with numerous records for New South Wales and Lord Howe Island (May, 1939; Lucas, 1914, 1935; Allender and Kraft, 1983; Lindauer et a/., 1961; Sonder, 1880); grows on rocky substrata in the subtidal to at least 25m, and common in kelp communities. Selected specimens examined: New South Wales: Coffs Harbour, 1-2m, 8-x-1985, P. and W. Farrant, UNSW 18623; Broughton I., 28-iv-1985, Farrant, UNSW 18029; Lord Howe I., Sep. 1908, Dun and Hedley, NSW 140443; Long Reef, 12-vii-1974, Harada R1269, NSW; Fairlight, 1-3m, 14-v-1985, Farrant, UNSW 18054; Dobroyd, 10m, 24-ix- 1985, Farrant, UNSW 18533; Jervis Bay, LWM, 18-11-1985, King, UNSW 17710. Queensland: Magnetic I., 13-v-1987, Phillips, MUCV 2568; Caloundra, Jan. 1909, Lucas, NSW 140438; Moreton Bay, Dec. 1913, Lucas, NSW 140437. Zonaria C. Agardh Growth from a marginal row of apical cells. Thalli with fronds in section having moderately distinct cortex and medulla, with medullary cells arranged in vertical tiers and cortical cells mostly paired to each medullary cell; sporangia with 8 aplanospores. There are three species of Zonaria in New South Wales, Z. diesingiana, Z. crenata and Z. angustata. The New South Wales specimens formerly referred to Zonaria sinclairit Hooker and Harvey are now placed in Homoeostrichus (see below). The taxon referred to as ‘Zonaria sp. in King and Farrant (1987) is Homoeostrichus olsentt. KEY TO THE SPECIES OF ZONARIA IN NEW SOUTH WALES iF Thallus narrow 2-4mm wide, linear, 10 cells thick, sporansdallsonllackingycellularmparaphyses)s sem mans ares ee ee eee Z. angustata 1* ‘Thallus lobed or fan-shaped (to 6cm wide), or linear with cuneate terminal segments (to 2cm wide), 6(-S)icells thick, sporangial/som, contaimime paraphyses= oo eee 2 ML, Thallus with lobed fronds somewhat denuded below, Sicellsithicks often covered by a networkjof hydnordier csc Z. crenata 2.* _Thallus lobed or broadly fan-shaped, not denuded below, (6-)Sicellsithickamnotcovered) by, hydroid eae ee etre ern Z. diesingiana Zonaria angustata (Keutzing) Papenfuss, 1952: 170. Womersley, 1987:248. Thalli 15-17cm long; fronds generally 2-4mm wide, more or less linear (Fig. 18A) but occasionally broader (to 20mm) in young plants or at the base; in section 10 cells thick (Fig. 18B); sporangial sori scattered on the distal frond surfaces, without cellular paraphyses. Seasonality: New South Wales collections of Zonara angustata made in January and August only. Australian distribution: from Elliston, South Australia, to Eden in New South Wales and around Tasmania; only two collections from New South Wales, from Eden and Green Cape, from the shallow sublittoral, 1-3m. Selected specimens examined: New South Wales: Eden, Jan. 1910, Lucas, NSW 140420; Green Cape, 1-3m, 17-viii-1987, D. May, MUCV 2571. Victoria: Newfield Bay PROC. LINN. SOC. N.SW., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 395 dis 5cem 9000 CRONE GbeDR80e & €300 008860 G0E00 eI es eal ents es ell esitea atest 6o8 50008 Soo FORA BBs S225 oooonDowo Smon Sood onesso5005 oOOOUDDOOVROWD jai=leleie} DOooOF7aGD ejelee\=)sse=stalSeis\S/5 oOOoOoDeSsS Oo GUBDSSOOOSB SS 90 000000855 Be 8 Oe eR Sep abos8s OF 98008 00 0 Becta bd bs bs bo0e Soa ee0o2eG 60 00 B 100um Fig. 18. Zonaria angustata: (A) habit (Eden, Jan. 1910, A. H. S. Lucas), (B) T.S. frond (MUCV 2571). (38°36’S, 142°51’E), 10-x11-1969, Womersley, NSW ex ADU A3408; Point Lonsdale, 4- x11-1985, Phillips, UNSW 19328. Zonaria crenata J. Agardh 1872:48, 1894:13. Womersley 1987:250. Thalli 5-15cm long; fronds 1-2cm wide, complanate, broader at the apex and usually denuded at the base (Fig. 19A), covered with a network of the hydroid Scoresbia diadala Watson (Fig. 19B,C); 6(-8) cells thick; sporangial sori scattered on the upper parts of the fronds. Plants of Z. crenata are often longer than those of Z diesingiana and have narrower and thinner fronds. They are often distinguished by the hydroid cover, though this feature alone is not sufficient for species identification. New South Wales specimens can PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 396 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Fig. 19. Zonaria crenata: (A) habit (MEL 16937), (B) habit of plant doubtfully distinct from Z. diesingiana but which is 6 cells thick and has hydroid cover (UNSW 17720), (C) frond apex. be difficult to place and there is considerable overlap between the two species with regard to characters such as frond thickness, frond shape and size, and the degree of hydroid cover. Some Z. crenata plants have a form similar to Z. diesingitna, with fronds 6 cells thick, complanate with broad tips and denuded lower axes and a cover of hydroid (Fig. 19B,C). Some Queensland specimens also have the typical Z. diesingiana form and yet have a cover of hydroid. The only typical Z. crenata specimens from New South Wales seen during this study were those collected from Sydney last century (MEL 16937). The relationship between the two taxa requires elaboration especially as Z. diesingiana is not recorded in southern Australia (Womersley, 1987). Seasonality: not determined. Australian distribution: from Fremantle, Western Australia, to Southport, Queensland, but rare in New South Wales; specimens examined by Womersley (1987) are either from the drift or from deep water. Selected specimens examined: New South Wales: Coffs Harbour, 29-vii-1980, Millar and Kraft, MELU AM 312 and MELU AM 313; Barrenjoey, 15m, 23-i11-1985, Farrant, UNSW 17720; Sydney, ex Herb. Sonder (1812-1881), MEL 16937; Plantation Point, 29-viii-1973, V. May, NSW; Green Point Warrah Sanctuary, 17-xii-1972, Larkum and Martin, NSW 140433. Queensland: Mooloolaba, 24-i-1944, V May 897, NSW. South Australia: Lefevre Peninsula, ex Herb. Sonder (1812-1881), MEL 16936, 16938; Glenelg, PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 39H, 22-viii-1946, V. May 2235, NSW; Kingston (36°50’S, 139°51’E), 16-x-1986, Womersley, NSW ex ADU A57343. a) @elaia|s):) 86 8O 8c 8 Bo ao eG 100um Fig. 20. Zonaria diesingiana: (A) habit of broad tall plant (UNSW 18019), (B) habit of plant with dense fronds (UNSW 18085), (C) UNSW 18292, (D) habit of broadly flabellate plant (UNSW 18022), (E) T:S. frond (UNSW 18085), (F) L.S. frond (UNSW 17755), (G) L.S. frond (UNSW 18008), (H) T.S. through sporangial sorus (UNSW 18006). Zonaria diesingiana J. Agardh, 1841:443; 1848:109. Allender and Kraft, 1983:77. Thalli (1.5-)4-11cm long, variable in habit, from erect to recumbent, often with a slight green or blue iridescence in sztu; fronds to 1.5-6cm wide, from linear to broad and flabellate (Fig. 20A-D), basal fronds often with a stupose central axis, but never completely bare of lateral membranous frond material; (6-)8 cells thick in mid-frond PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 398 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES regions (Fig. 20E-G); sporangial plants with sori arranged irregularly or in roughly con- centric lines, and with sterile paraphyses (Fig. 20H). There are problems in distinguishing Zonaria diesingiana from Z. crenata in New South Wales (see notes under that species). Zonaria turnertana has been reported widely from the New South Wales coast (Womersley, 1967; May et al., 1970; May and Larkum, 1981). These records are here recognized as of Z. diesingiana. Seasonality: sporangial plants collected in all months of the year. Australian distribution: reported from Sydney to Coffs Harbour in New South Wales and Lord Howe Island (Allender and Kraft, 1983); also at Norfolk Island and on the New South Wales coast south to Green Cape; the most abundant dictyotalean alga on the central New South Wales coast (King and Farrant, 1987), at Lord Howe Island (Allender and Kraft, 1983) and in the Coffs Harbour region (A. Millar, pers. comm.), in both sheltered and open coast localities in eulittoral rock pools and to depths of 20m; abundance declines markedly with distance south and north of the central New South Wales region. Selected specimens examined: New South Wales: Hastings Point, 19-v-1986, Phillips, MUCV 2561; Coffs Harbour, 1-2m, 8-x-1985, Farrant, UNSW 18618; Diamond Head, 0-1m, 10-x-1985, PR and W. Farrant, UNSW 18669; Fairlight, 1-2m, 26-xi-1985, Farrant, UNSW 18582; Fairlight, 5m, 29-iv-1986, Farrant, UNSW 18846; Kiama, July 1899, Lucas, NSW; Jervis Bay, LWM, 18-11-1985, King, UNSW 17711; Broulee, 21-viii- 1987, D. May, MUCV 2574; Malau Bay, 21-viii-1987, D) May, MUCV 2569; Barragga Point, LWM, 13-xi-1986, Farrant, UNSW 19360. Queensland: Noosa, 4-x11-1976, King and Kertesz, UNSW 15313. COMPARISON WITH ZONARIA SPECIES IN ADJACENT REGIONS Four species are found in southern Australia, Zonaria angustata, Z. crenata, Z. spiralis (J. Agardh) Papenfuss and Z. turneriana J. Agardh. Zonaria spiralis has a southern distri- bution from Rottnest Island, Western Australia, to Flinders, Victoria, and has charac- teristic spirally-twisted branches. Zonaria turneriana is found from Geraldton, Western Australia, to Port Phillip Heads, Victoria, Tasmania, and New Zealand. This species has mature fronds that are more or less linear, and 8-10 cells thick, but which are broader than those of Z. angustata (2-5[-8]mm). Only Z. crenata and Z. diesingiana occur in Queensland. Homoeostrichus J. Agardh Growth from a marginal row of apical cells. Thalli in transverse section having cells of medulla of uniform size and arranged in regular tiers, cells of the cortex mostly unpaired; sporangia with 4 aplanospores. The genus Homoeostrichus was formerly placed within Zonaria (Papenfuss, 1944), but has been recently re-established by Womersley (1987). Homoeostrichus can be distin- guished from Zonaria by sporangia with 4 aplanospores, as yet only observed in H. sinclairit and H. olsen (cf. 8 in Zonaria), and by mostly unpaired cortical cells (which are mostly paired in Zonaria). There are two species of Homoeostrichus in New South Wales, H. olsen and H. sinclair; both can be difficult to separate from Zonaria diesingiana. KEY TO THE SPECIES OF HOMOEOSTRICHUS IN NEW SOUTH WALES ie Thallus broadly fan-shaped, greyish, 5(-6) cells thick belowsapexn.2a sh) Paitin iv Bias ee see EN aie tose FH. olsenu PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 399 1.* Thallus with cuneate (wedge-shaped) terminal segments, light brown, denuded below, 6 cells thick except thick ergatysaal aria eye) seccnt Senne ON At he UMMeRe cis! soc We Urnhet, tale Ge) ate: nies Bt aya HZ. sinclairi Ri KAN sii Pao st Tou jeane SDoooo Ir a see eee ee eesieeseven 100um Fig. 21. Homoeostrichus olsenii: (A) habit of sporangial plant (UNSW 18242), (B) habit of male plant (UNSW 18243), (C) section of sporangial plant (UNSW 18216), (D) section of male plant (UNSW 18243). Homoeostrichus olsenit Womersley, 1987:243. Thalli 9-12cm long, a greyish colour; fronds 1-3(-6)cm wide (Fig. 21A,B), com- planate, flabellate or irregular, with dense hairs covering most of the under surface; 5(-6) cells thick (Fig. 21C,D); sporangial sori (Fig. 21C) scattered across the frond in roughtly concentric rows; male gametangia in orange-brown sori without paraphyses (Fig. 21D). Seasonality: sporangial plants collected in April, July, September, October, and November; male plants in April and July. Australian distribution: south-eastern Australia, Nora Creina, South Australia, to Sydney Harbour (Womersley, 1987); in the present study from three relatively rough water localities in the Sydney region, to depths of 21m. Selected specimens examined: New South Wales: Fairlight, 2-3m, 16-ix-1985, Farrant, UNSW 18293; Fairlight, 2-3m, 8-vii-1985, Farrant, UNSW 18216; Dobroyd, 8-iv- 1979, Farrant, UNSW 18779; Bare I., 6m, 25-vi1-1985, Farrant, UNSW 18242 and UNSW 18243; Bare I., 10m, 17-x-1985, Farrant, UNSW 18567; Boat Harbour, 21m, 21-iv-1975, Harada R2361, NSW. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 400 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Fig. 22. Homoeostrichus sinclairu: (A) habit (UNSW 18020), (B) T.S. through frond ‘midrib (UNSW 18004). Homoeostrichus sinclair (Hooker and Harvey) J. Agardh, 1894:15; Womersley, 1987:242. Zonaria sinclairtt Hooker and Harvey, 1845:530. Thalli 8-18cm long; fronds linear with cuneate terminal segments to about lcm wide, narrowing below to a bare ‘midrib (Fig. 22A); hairs largely on ‘midrib’; in trans- verse section generally 6 cells thick below the apex and expanded at the ‘midrib’ (Fig. 22B); sporangia with multicellular pedicels, amongst hairs. Seasonality: collected throughout the year, with sporangial plants in all except the summer months; gametangial plants unknown for this species. Australian distribution: southern and eastern Australia (Great Australian Bight to Newcastle) (Womersley, 1987), and as far north as Broughton Island, in relatively rough water localities and at depths greater than 2m. Selected specimens examined: New South Wales: Broughton I., 2m, 28-iv-1985, Farrant, UNSW 18027; Norah Head, 19-iv-1978, Kertesz, UNSW 15308; Long Reef, 2- PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P.A. FARRANT ANDR. J. KING 401 vii-1975, Harada R4024, NSW; Dobroyd, 10m, 24-1x-1985, Farrant, UNSW 18535; Bare I., 5m, 6-11-1986, Farrant, UNSW 18301; Plantation Point, 26-i1-1974, Larkum, NSW. COMPARISON WITH HOMOEOSTRICHUS SPECIES IN ADJACENT REGIONS A third species, Homoeostrichus canaliculatus (J. Agardh) J. Agardh, is found in southern Australia but not in New South Wales. This species has linear fronds mostly 1- 1.5mm wide and 6-7 cells thick (Womersley, 1987). Stypopodium Kuetzing Growth from a marginal row of apical cells. Thalli prostrate or erect, membran- ous; fronds in transverse section having cortical cells smaller than medullary cells and medullary cells not strictly tiered in transverse section; no sterile paraphyses associated with sporangia. Stypopodium can easily be confused with Yaonza. It differs from Taonia in having thick distal fronds, the fronds becoming 4 cells thick close to the apex (Zaonza has thin distal fronds and becomes 4 cells thick at a distance from the apex, Fig. 23A); in Stypopodium there is an abrupt change in the size of cells from the cortex to the medulla with pig- mented cortical cells much smaller (in Zaonza there 1s no such abrupt change in cell size, Fig. 23B). Allender and Kraft (1983) also refer to the smoothly arching laciniae in most clumps of Taonza which are not seen in S. australasica (Zanardini) Allender and Kraft. Stypopodium flabelliforme is the only species of the genus from the mainland New South Wales coast, although specimens exhibit some gradation towards both S. aus- tralasicum and S. flabelliforme var. rabdoides Allender and Kraft, both recorded for Lord Howe Island by Allender and Kraft (1983). Mainland New South Wales specimens are variable in habit (erect with single holdfast to decumbent with overlapping blades), frond shape (flabellate to lobed), and colour (light brown, iridescent, or with slightly darker longitudinal bands). Stypopodium flabelliforme Weber-van Bosse, 1913:176. Allender and Kraft, 1983: 96. Thalli 4-8cm long, usually decumbent although larger plants erect, brown, often iridescent (blue-green) with dark longitudinal bands not obvious on the brown dried thalli; fronds 2-9cm wide, flabellate and sometimes divided into irregular lobes (Fig. 24A,B); regular concentric hair lines sometimes present, especially on larger flabellate plants (Fig. 24A); fronds in section having a medulla 4-6 cells thick in the lower parts (Fig. 24C,D); sporangia densely scattered all over the fronds (except for conspicuous sterile bands next to the hair lines), or in patches. Seasonality: collected in most months, sporangial plants in February, April and October; no gametangial plants. Australian distribution: Lord Howe Island and the southern Great Barrier Reef (Allender and Kraft, 1983); Allender and Kraft (1983) were the first to record the species for the Australian mainland coast (Coffs Harbour, New South Wales, and Heron I., Queensland); in this study collected south to Jervis Bay, from the upper sublittoral to depths of 20m, mostly on open coasts. Selected specimens examined: New South Wales: NW Solitary I., 10-16m, 6-x- 1985, Farrant, UNSW 18603; Seal Rocks, 0-2m, 10-x-1985, P and W. Farrant, UNSW 18679; Clovelly, 1-3m, 16-iv-1985, Farrant, UNSW 17781; Jervis Bay, 15m, 14-iv-1984, Farrant, UNSW 16201; Plantation Point, 30-vii-1973, V. May, NSW; Malau Bay, 21-vii- 1987, D. May, MUCV 2570. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 402 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES If Ss fy GEOCGSOSSS. A B fa) sa aaa bode We ae Sy cae ae Oe ese ea ce ao, ipo SE ck pi Fig. 23 (above). Diagrams showing the differences between Taonia (T) and Stypopodium (S): (A) L.S. apex, (B) T.S. lower part of frond. Fig. 24 (below). Stypopodium flabelliforme: (A) habit of broad flabellate plant (UNSW 18601), (B) habit of smaller more prostrate plant (UNSW 17770), (C) L.S. through apex (UNSW 18043), (D) L.S. through frond (UNSW 18014). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 403 COMPARISON WITH STYPOPODIUM SPECIES IN ADJACENT REGIONS Two other taxa of Stypopodium are found at Lord Howe Island, S. flabelliforme var. rabdoides and S. australasicum (Allender and Kraft, 1983), and a further two have been recorded for northern Australia, S. zonale (Lamouroux) Papenfuss and S. lobatum Kuetz- ing (Lewis, 1985). The genus is not recorded in southern Australia. Taonia J. Agardh Growth from a marginal row of apical cells. Thalli erect, membranous; fronds flabellate, often deeply dissected; surface cells similar in size to medullary cells, distal part of frond thin, two cells thick, thicker in lower parts; sori scattered on frond in more or less concentric zones. Taonia can be easily confused with Stypopodium (Fig. 23A,B) (see notes under that species). There is only one species of Taonza in New South Wales, T? australasica. Us OCD poceececcnans pee erceesseee = Ci me a [3 ere : 3 ayIcn aaeese OG DOOOOOUBOOUOOB SS OOD0O000 ES Pa aaases P2989 F 200um Fig. 25. Taonia australasica: (A) habit of large, broad plant (UNSW 18070), (B) habit of narrow fronded plant (UNSW 17634), (C) habit of broad plant (UNSW 17774), (D) T.S. through sporangium (UNSW 17634), (E) L.S. through apex (UNSW 18066), (F) T.S. through male gametangial sorus (UNSW 18234). PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 404 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES Taonia australasica J. Agardh, 1894:30. Allender and Kraft, 1983:91; Womersley, 1987:238. Thalli 4-19cm long, tan-olive in colour at the top, dark brown at the base; fronds 0.5-17cm wide, flabellate (Fig. 25A-C), in transverse section cells similar in size through- out (Fig. 25D), thin, remaining 2 cells thick for some distance below the apex (Fig. 25E); sporangia borne on two stalk cells at maturity, sessile or partially embedded in the thallus (Fig. 25D); antheridia in indusiate sori on male plants (Fig. 25F). Seasonality: collected in all months except December, but seasonally abundant with marked decrease in abundance and biomass in summer (King and Farrant, 1987); sporangial plants collected in all months of the year except October and December; the single male gametangial plant collected in July the first record of sexual reproduction for this species. Australian distribution: south-eastern Australia, from Kangaroo Island, South Australia, to Coffs Harbour, New South Wales (Womersley, 1987), and Lord Howe Island (Allender and Kraft, 1983); found in the sublittoral below 2m depth at both open coast and estuarine localities (Sydney Harbour). Selected specimens examined: New South Wales: Broughton I., 2m, 28-iv-1985, Farrant, UNSW 18024; Harbord, 1-2m, 16-11-1985, Farrant, UNSW 17610; Fairlight, 1- 2m, 16-iv-1986, Farrant, UNSW 18358; Mrs Macquarie’s Point, 2m, 27-vi-1985, Farrant and Puttock, UNSW 18211. COMPARISON WITH TAONIA SPECIES IN ADJACENT REGIONS Taonia australasica is the only species of Taonia found in Australia. It is found in the regions adjacent to New South Wales, at Lord Howe Island, in Queensland and in southern Australia. CONCLUSIONS The algae of the New South Wales coastline have been much less studied than those on adjacent shores, especially those to the south. The Dictyotales are well represented in the flora with 22 species in 13 genera, and since they are abundant during most seasons especially in the upper sublittoral (King and Farrant, 1987) they are frequently referred to in ecological accounts. The present treatment has clarified the species encounted in New South Wales and provides keys and descriptions for their ready identification. ACKNOWLEDGEMENTS We would like to thank E. O’Brien, W. Farrant, and C. F. Puttock, for valuable assistance with field work; Dr B. M. Allender, Dr G. Kraft, D. May, Dr A. J. K. Millar, Dr J. Phillips, Mrs D. Sinkora, and Prof. H. B. S. Womersley for loan of specimens and for helpful discussion during the study; and MEL, MUCV and MELU herbaria for loan of specimens. References AGARDH, J. G., 1841. — In historiam algarum symbolae. Linnaea 15: 1-50. , 1848., — Species, Genera et Ordines Algarum. Vol. 1. Lund: Gleerup. ——, 1872. — Tillalgernes systematik. Acta Univ. Lund. 9: 1-71. ——, 1882. — Tillalgernes systematik. Acta Univ. Lund. 17: 1-136, pls 1-3. ——, 1894. — Analecta Algologica. Cont. I. Acta Univ. Lund. 29: 1-144, pl. 1. ALLENDER, B. M., 1980. — Dictyotopsis propagulifera (Phaeophyta) — an algal enigma. Phycologia 19: 234-236. ,and Krart, G. T., 1983. — The marine algae of Lord Howe Island (New South Wales): the Dictyotales and Cutleriales (Phaeophyta). Brunonia 6: 73-130. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 P. A. FARRANT ANDR. J. KING 405 ARESCHOUG, J. E., 1854. — Phyceae novae et minus cognitae in maribus extraeuropaeis collectae. Acta Reg. Soc. Sct. Upsala, Ser IIT 1: 329-372. BOERGESEN, F., 1935. — A list of marine algae from Bombay. K. Dan. Vidensk. Selsk. Biol. Meddr 12: 1-64, pls. 1-10. BOLD, H. C., and WYNNE, M. J., 1985. — Introduction to the Algae. Structure and Reproduction. 2nd Edition. Englewood Cliff, New Jersey: Prentice-Hall Inc. BOROWITKZA, M. A., KING, R. J., and LARKuM, A. W. D., 1982. — Field guide to the marine plants of New South Wales. Chlorophyta, Phaeophyta, and Seagrasses. (Privately publ., Sydney). 70 pp. Bory DE SAINT-VINCENT, J. B., 1827. — Padine. Duct. Class. Hist. Nat. 12: 589-91. Paris. Cribs, A. B., 1954. — Records of marine algae from south-eastern Queensland, I. Pap. Dep. Bot. Univ. Qd 3: 15-37. EARLE, S. A., 1969. — Phaeophyta of the eastern Gulf of Mexico. Phycologia 7: 71-254. EDELSTEIN, T., and WOMERSLEY, H. B. S., 1975. — The thallus and spore development of Lobospira bicus- pidata Areschoug (Dictyotales): Phaeophyta. Trans Roy. Soc. S. Aust. 99: 149-156. Gepp, A., and GeppP, E. S., 1906. — Some marine algae from New South Wales. J. Bot. 44: 249-261. GREVILLE, R. K., 1830. — Algae Britannicae. Edinburgh: MacLachlan & Stewart. Harvey, W. H., 1860. — Phycologia Australica. Vol. III1., London: Reeve. Hauck, F., 1887. — Uber einige von J. M. Hildebrandt in Rothen Meere und Indischen Ocean gesammelte Algen. Hedwigia 2: 41-45. Haupt, A. W., 1932. — Structure and development of Zonaria farlowi. Am. J. Bot. 19: 239-254, pls XIX-XXII. HOOKER, J. D., and Harvey, W. H., 1845. — Algae Novae Zelandiae. Lond. J. Bot. 4: 521-551. KING, R. J., and FARRANT, P. A., 1987. — The phenology of the Dictyotales (Phaeophyceae) at a sheltered locality in Sydney Harbour, New South Wales, Australia. Bot. Marina 30: 341-350. LAMOUROUX, J. V. F., 1809. — Observations sur la Physiologie des Algues marines, et description de cing nouveaux genres de cette famille. Nouv. Bull. Sci. Soc. Philom. Paris 1: 330-333, pl. 6. LEvVRING, T., 1940. — Die Phaeophyceengattungen Chlanodiphora, Distromium, und Syringoderma. K. fyswogr. sallsk. Lund. Forh. 10: 1-11. Lewis, J. A., 1985. — Checklist and bibliography of benthic marine macroalgae recorded from northern Australia. II. Phaeophyta. Department of Defence Materials Research Laboratories, Victoria. Report MRL-R-962, 40 pp. LINDAUER, V. W., CHAPMAN, V. J., and AIKEN, M., 1961. — The marine algae of New Zealand. II: Phaeophyceae. Nova Hedwigia 3: 129-350. Lucas, A. H. S., 1909. — Revised list of the Fucoideae and Florideae of Australia. Proc. Linn. Soc. N.S.W. 34: 9-61. ——., 1913. — Notes on Australian marine algae. 1. Fucoideae. Proc. Linn. Soc. N.S.W. 38: 49-60. ——., 1914. — Marine algae. British Association for the Advancement of Science. Handbook for N.S.W.: 459-463. Sydney: Edward Lee & Co. ——,, 1935. — The marine algae of Lord Howe Island. Proc. Linn. Soc. N.S.W. 60: 194-232. May, V., 1939. — A key to the marine algae of New South Wales — Part II. Melanophyceae (Phaeophyceae). Proc. Linn. Soc. N.S.W. 64: 191-215. ——.,, BENNETT, I., and THOMPSON, T. E., 1970. — Herbivore-algae relationships on a coastal rock platform (Cape Banks, N.S.W.). Oecologia (Berl. ) 6: 1-14. ——., COLLINS, A. J., and COLLETT, L. C., 1978. — A comparative study of epiphytic algal communities on two common genera of sea-grasses in eastern Australia. Aust. J. Ecol. 3: 91-104. , and LARKuM, A. W. D., 1981. — A subtidal transect in Jervis Bay, New South Wales. Aust. J. Ecol. 6: 439-457, NGAN, Y., and Price, I. R., 1979. — The intertidal algae of the mainland coast in the vicinity of Townsville, Queensland, tropical Australia. Atoll Res. Bull. 237: 1-29. , and , 1980. — Seasonal growth and reproduction of intertidal algae in the Townsville region (Queensland, Australia). Ag. Bot. 9: 117-134. PAPENFUSS, G. F., 1943. — Notes on algal nomenclature. II. Gymnosorus J. Agardh. Am. J. Bot. 30: 463-468. , 1944. — Notes on algal nomenclature. III. Miscellaneous species of Chlorophyceae, Phaeophyceae and Rhodophyceae. Farlowza 1: 337-346. ——.,, 1952. — Notes on South African marine algae. III. J. S. Afr. Bot. 17: 167-188. READERS DIGEST, 1977. — Atlas of Australia. Sydney: Reader’s Digest Services Pty Ltd. SONDER, W., 1871. — Die Algen des tropischen Australiens. AbA. naturw. Ver. Hamburg 5: 33-74, plates 1-6. SONDER, O. G., 1880. — I. Algae Australianae hactenus cognitae. In: Fragmenta phytographiae Australiae, Suppl. ad. Vol. II (F. Mueller): 1-42. Melbourne. WEBERYAN BossE, A., 1913. — Liste des algues du Siboga. I. Myxophyceae, Chlorophyceae, Phaeophyceae. Stboga Exped. Monogr. 59a: 1-186. PROC. LINN. SOC. N.S, 110 (4), (1988) 1989 406 THE DICTYOTALES (ALGAE PHAEOPHYTA) OF NEW SOUTH WALES WOMERSLEY, H. B. S., 1949. — Studies on the marine algae of southern Australia. No. 3. Notes on Dicty- opteris Lamouroux. Trans Roy. Soc. S. Aust. 73: 113-116. ——, 1967. — A critical survey of the marine algae of southern Australia. II. Phaeophyta. Aust. J. Bot. 15: 189-270. —, 1981. — Biogeography of Australasian marine macroalgae. In: M. N. CLAYTON and R. J. KING, (eds), Marine Botany: an Australasian Perspective: 292-307. Melbourne: Longman Cheshire. ——., 1987. — The Marine Benthic Flora of Southern Australia. Part II. Adelaide: South Australian Government Printing Division. , and BAILEY, A., 1970. — Marine algae of the Solomon Islands. Phil. Trans Roy. Soc. Lond. B. Biol. Scz. 2592-32: YAMADA, Y. 1931. — Notes on some Japanese algae. II. J. Fac. Sci. Hokkaido Imp. Univ. Ser. 5 1: 65-76. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 A distinctive new Species and new Distribution Records of Stilbopteryginae (Insecta: Neuroptera: Myrmeleontidae) C. N. SMITHERS SMITHERS, C. N. A distinctive new species and new distribution records of Stilboptery- ginae (Insecta: Neuroptera: Myrmeleontidae). Proc. Linn. Soc. N.S.W. 110 (4), (1988) 1989: 407-410. Stilbopteryx mouldsorum sp. n., a very distinctive species of lacewing, is described from Western Australia. It has a colour pattern not found in any known species of the genus. New distribution records, based on material in the collections of the Australian and Queensland Museums are given for the other species of Stilbopteryginae. C. N. Smithers, Research Associate, Australian Museum, College Street, Sydney, Australia 2000; manuscript received 12 April 1988, accepted for publication 20 July 1988. INTRODUCTION The genus Stz/bopteryx Newman, with six described species, and Aeropteryx Riek, with three species, constitute the subfamily Stilbopteryginae of the Myrmeleontidae. They include some of the largest and most spectacular Australian lacewings. Some species have a wing length of up to 50mm. Very little is known of the biology of this group. Of only one species (S. linearis Navas) is the larva known (McFarland, 1968). Development from egg to adult took six years in culture. Both sexes of the species of Stil- bopteryx are known, except for S. albosetosa Riek of which males have not been found. All three species of Aeropteryx are known from males only. Riek (1968) published a key which included six of the seven species known at the time. Later (Riek, 1976) he described two more species and included them, with all seven previously known, in a revised key. He also listed the synonyms for each and gave information on the distribution of the species. New (1982) discussed the status of the genera, which at that time were usually included in the family Stilbopterygidae. He gave reasons for considering them to constitute a subfamily of the Myrmeleontidae. During study of collections in the Australian and Queensland Museums a female of a very distinctive, undescribed species of Stzlbopteryx was found as well as material which increased knowledge of the distribution of the other species. This material forms the basis of this note. For the sake of completeness the previously known distributions of the species are summarized. In the list of material the initials ‘M.S.M? and ‘B.J.M’ indicate specimens collected by M. S. and B. J. Moulds respectively and ‘AM’ and ‘QM’ the Australian and Queens- land Museums. DESCRIPTION AND RECORDS Stilbopteryx mouldsorum sp.n. Female: Head very dark brown, slightly paler on each side of epicranial suture. Front of head anterior to lower level of eyes creamy yellow. Antennae very dark brown, club with pale transverse bands. Setae on scape black. Pronotum dark brown with creamy yellow bands along anterior and posterior margins. Setae black. Pterothorax dorsally dark brown. Setae on anterior half of mesonotum mostly black, some thoracic setae white. Thorax laterally grey-brown, setae mostly white. Legs very dark brown. Abdominal PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 408 A NEW LACEWING segment 1 very dark brown; segment 2 very dark brown with narrow creamy yellow posterior border; segments 3-6 very dark brown with very broad anterior and posterior creamy yellow bands; segment 7 similar but bands narrower. There is as much creamy yellow area as darker colour in the basal half of the abdomen. Ectoprocts very dark brown with creamy yellow postero-dorsal border and similarly coloured patch near base. Eighth and ninth tergites very dark brown with paler hind borders. Posterior gonapophyses long and narrow with black setae, similar to those of S. linearis Navas. Fore wings with costal area hyaline but dark brown just basad of creamy yellow pterostigma. Radial area with infuscation reaching hind margin of area distally before pterostigma. The dark brown band formed by these coloured areas is continued beyond the pterostigma adjacent to the wing margin anteriorly, around the wing apex and along hind margin of wing, tapering to end about half way along hind margin. Costal cross- veins, Rs and its branches and veins and crossveins posterior to CuA creamy yellow. Other veins dark brown. Hind wing costal area brown almost from base with crossveins nearly all brown. Other markings as in fore wing but veins and crossveins posterior to CuA mostly brown. Fore wing length: 45mm. Body length: 42mm. Male: Not known. Material examined: | 9 (holotype), Western Australia, 3km from West Strelley River, approx. 70km SE Port Hedland, 16.11.1977. M.S. and B.J. Moulds. Holotype in Aus- tralian Museum. Discussion: Stilbopteryx mouldsorum is recognizable by its colour pattern. It is the only species in the subfamily in which the brown band along the front of the wing is con- tinued beyond the apex and along the hind margin. The creamy yellow pronotal marks are more extensive and stand out more clearly than in other species. The similarly coloured areas on the abdomen are so extensive and conspicuous that they give the abdomen a banded appearance. In other species these areas are much smaller and appear as isolated spots. Stilbopteryx linearis Navas Recorded distribution. South Australia)’ SE Lake Eyre North, near Muloorina homestead; Prescott Point, Madigan Gulf, Lake Eyre North; Killalpanima, 160km E Lake Eyre. New record. South Australia: 1 9, Arkoola, northern Flinders Ranges, 21.1.1976, M.S.M. and B.J.M. (AM). Stilbopteryx albosetosa Riek Recorded distribution. Northern Territory; Murella Park; Daly Waters; Jabaluka Lagoon, 14km N. Mudginbarry head station; 1km S Cahill’s Crossing, East Alligator R; 5km NNW Cahill’s Crossing, East Alligator R. New records. Northern Territory: 3 9 , Waterhouse R., Mataranka Hstd., 23.x11.1968; 19, McArthur R., 15km S junction Carpentaria/Tablelands Highway, 12.1.1986, M.S.M. and B.J.M. (AM). Queensland: 19, Almaden, Chillagoe dist., 1.1932, W. D. Campbell; 19, swamp, 28km N Leura, 30.xi.1974; 29, East Haydon, 60km SE Normanton, 15.1.1986, M.S.M. and B.J.M. (AM). Stilbopteryx walkert Kimmins Recorded distribution. Queensland: Moreton Bay; Fletcher; Eidsvold; Rifle Range, Biggenden; Bundaberg; Crystal Creek, 35km SE Ingham; Clermont; Stradbroke Is.; Carnarvon Range; Almaden, Chillagoe dist. New records. Queensland: 20°, Moffat Nat. Park, The Chimneys, 14.x11.1987, Monteith, PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 C. N. SMITHERS 409 Thompson, Yeates; 1 9, Blackdown Tablelands, via Dingo, 1984, S Pearson (QM); 19, Miriam Vale, Mt Larcom area, 27.xu1.1966; 19, Bauple, 14.x11.1977; 10°, Doolandella, Brisbane, 12.1.1985; 10°, Boomerang Range, 30km W Marlborough, 8.xi1.1980; 19, Lawn Hill Creek, Adel’s Grove, W Gregory Downs, 19.x11.1986, M.S.M. and B.J.M. (AM). New South Wales: 19, 20km N Grafton, 21.1.1975, J. Frazier; 10°, Calumet, 26km NE Binnaway, 16.xii.1931, C. F. Garnsey; 19, Wheeogo, near Dunedoo, 1.1971, W. Gaden; 19, Mann R., 5km upstream from Old Glen Innes—Grafton Rd., crossing, 16.x1.1977, M.S.M. and B.J.M. (AM). Stilbopteryx napoleo (Lefebvre) Recorded distribution. Western Australia: Waroona; Yallingup; Swan R.; Yanchep; Perth; ‘NW New Holland’; ‘South and west Autralia. Northern Territory: 99km NNW Alice Springs. South Australia: Coorong Nat. Park. New records. Western Australia: 19, 30km S Mendurah, 13.1.1971; 19, Waylunga Nat. Park, 35km NW Perth, 6.1.1971; 20°, 19, John Forrest Nat. Park, Darling Ranges, 21- 23.1.1971, G. A. Holloway and H. Hughes (AM). Stilbopteryx costalis Newman Recorded distribution. New South Wales: Sydney; Shoalhaven; 42km NE Binnaway. Queensland: Stradbroke Is. New records. Queensland: 10°, Brisbane, 4.xi.1936; 10°, 19, Carnarvon Range, 7.xi1.1941; 19, Milmerran, 21.xii.1941, J. Macqueen (QM); 19, Cloncurry, 19.1.1984, M.S.M. and B.J.M. (AM). New South Wales: 19, Waterfall, 31.x11.1978, D. Doolan; 10, Loftus, 27.xii.1978, J. Olive; 10°, Engadine, 16. xi1.1976, M. Schneider; 10°, Como, 9.xi.1961, L. Willan; 1 9, Norton’s Basin, Nepean R., 19.x11.1916, A. Musgrave (AM). Stilbopteryx auricornis Kimmins Recorded distribution. Western Australia’ Sherlock R.; Pinjara; 32km WSW Mt Magnet; 30km SE by S Carnarvon. New records. Western Australia: 19, Carnarvon, 25.11.1977, M.S.M. and B.J.M. 19, Marble Bar, 18.1.1974, G. Jones (AM). South Australia: 10°, 1 9, Stuart Highway, 56km S Northern Territory border, 4.11.1986, M.S.M. and B.J.M. (AM). Northern Territory: 39 , Yalora Resort, Ayer’s Rock, 1.11.1984, M.S.M. and B.J.M. (AM). Aeropteryx brocki (Manski) Recorded distribution. Queensland: Kalpower; Little Crystal Creek, Mt Spec; Bluff Range, Biggenden. New records. Queensland: 10, Edungalba, near Duaringa, 24.1.1982, M.S.M. and B.J.M. (AM); 20°, Blackdown ‘Tableland, via Dingo, xi1.1978, Maywald, Czechura (QM). Aeropteryx gibba Riek Recorded distribution. Northern Territory: Deaf Adder Creek; Murella Park; Katherine Gorge; 7km NW by N Cahill’s Crossing, East Alligator R.; 9km N by E Mudginbarry head station. New records. Northern Territory; 10, Barkly Highway, 75km ESE junction with Tablelands Highway, 4.1,1987. M.S.M. and B.J.M. (AM). Queensland: 30°, Little R., E Croydon, 15.xii.1986; 10°, Iron Range, x1.1985, G. Wood (QM). Western Australia: 20°, Kununurra, 6-7.1.1986, M.S.M. and B.J.M. (AM). PROC. LINN. SOC. N.SW,, 110 (4), (1988) 1989 410 A NEW LACEWING Aeropteryx monstrosa Riek Recorded distribution. Queensland: Station Creek Spring, Silver Plains, Cape York. Eidsvold. Hammond Is. New record. Queensland: 10°, Station Creek, 15km N Mt Molloy, 30.xii.1973, A. and M. Walford-Huggins (AM). ACKNOWLEDGEMENTS I would like to thank the many collectors of the material listed, especially Max and Barbara Moulds, for the considerable amount of material donated to the Australian Museum. The Director of the Queensland Museum kindly allowed me to study its material. References MCFARLAND, N., 1968. — (Cover picture.) Friends of S. Aust. Mus. 7: 1-2. New, T. R., 1982. — A reappraisal of the status of the Stilbopterygidae. (Neuroptera: Myrmeleontidae). /. Aust. ent. Soc. 21: 71-75. RIEK, E. F., 1968. — A new genus and key to species of Australian Stilbopterygidae (Neuroptera). J. Aust. ent. Soc. 7; 105-108. ———, 1976. — The family Stilbopterygidae (Neuroptera) in Australia. J. Aust. ent. Soc. 15: 297-302. PROC. LINN. SOC. N.S.W., 110 (4), (1988) 1989 Albludomelitawa mOtenaseuomcns tence) ele Adamson, D., see Glasby, P. Index VOLUME 110 Page 337 @ladoce rae sass ea a eee eee Acarina: Erythraecidae ................ 193 Additional observations on WNitella verti- cillata (Characeae) from a new locality in New South Wales, by A. T. Hotchkiss Sep oimalO rie ee cepvce ris ssekcoge steve suse 187 Albertson, E. L., see Rowe, F. W. E. Aloaesbhacophy tay. 61s asta cies at 369 Ninja apex) sseccudecdeboubo Boop aan oS 327 Arachnida: Pseudoscorpionida ......... 289 Archer, A. W., The lichen genus Cladonia section Cocciferaein Australia ........ 205 XS tenold Cale qe sinc ae aleve ictcalta halt 83 Associations of some Glycaspis species (Homoptera: Spondyliaspididae) with their Eucalyptus species hosts, by K. M. IO ORC Mansa heute gcuteycu ena tase tease cs 19 Barraba, Late Devonian trilobites from . . 27 Baseline survey of the benthic macrofauna of Twofold Bay, N.S.W., with a dis- cussion of the marine species intro- duced into the bay, by P. A. Hutchings, J. T. van der Velde & S. J. Keable ..... 339 Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western Blue Moun- tains, by P. Glasby, P. M. Selkirk, D. Adamson, A. J. Downing & D. R. Sel kicker cpa taes hove otto oashers te eugesuses 141 Botany Bay, ichthyoplankton ........... 1 Brewer, D. T., & Warburton, K., A dietary study of Szllago analis and its variation in three Australian localities ........... 215 Galandrniayseedity pes aa ae eyes eee 307 Giallanoidayie sees skea\shersyyensiypens ushers eins 297 Callidosomatine mite ................. 193 Campbell, K. G. & Moore, K. M., Two new species of Glycaspis (Homoptera: Spondyliaspididae) from _ potentially endangered Eucalyptus species and one from E. stricta....... Sisha MaMa OAR are 11 Capilo, a new group of subgen. Neurochaeta s. stra l(Diptera) yee. teeicusy weg doe re ek 46 Carastrum ferrari, gen. et sp. Nov. ......... 195 Carolin, R. C., see Syeda, S. T. Carr, S. G. M., & Carr, D. J., Sir William Macleay Memorial Lecture 1987. The elastic-sided gumleaf, or: the rubber cuticle and other studies of the Corymbo- SA oe el nner hes icy at wuss ear es matter rte st ie 101 @araceae ha Maria sl Meale taille cue ei 175, 187 Charletoniini, mite tribe now defined .... 203 Cladonia sect. Cocciferae (lichens) ......... Gladontatancustatamyaer tre trae oe Gladontatbimbentensiswne nemo ore Gladonianflocrkanapew aetna ere ee Gladontaimacilentampe eee Gladontaimuntay eee one Gladoniarbleuroiai eee ee ea eee: Gladontarsubdievtatajers ier yee eee Gladonia weymouthid se). haa i Corymbosae, rubber cuticle .............. Crustacea, Amphipoda Crustacea, ostracodes ....5..).........- Cyclopoida Devonianitmlobites jae ee see Dictyopterts acrostichoides .......-........ Diciyotavaltemmiidapae aera Dictyotalbartayrestty weno sia ee eee Dictyotales (Algae: Phaeophyta) of New South Wales, by P. A. Farrant & R. J. RT eek ch asralierean wea ees At ata Dietary study of Sillago analis and its vari- ation in three Australian localities, by D. T. Brewer & K. Warburton ....... Dilophussintenmedtusse a se oy ee Dilophuswmargrnatusy ys.) serie ee Diptera: Neurochaetidae ............... Distinctive new species and new distri- bution records of Stilbopteryginae (In- secta: Neuroptera: Myrmeleontidae), byg@aNWSmnuthersiyyaria ese creer Distribution and ecology of Recent ostra- codes (Crustacea) from Port Hacking, New South Wales, by I. Yassini & A. J. Wright Distromium flabellatum .......-0.0-0205- Diurnal survey of ichthyoplankton abun- dance, distribution and seasonality in Botany Bay, New South Wales, by A. S. Stefferé Ba Cabeasemeinri ee aes Dodson, J. R., see Kodela, P. G. Downing, A. J., see Glasby, P. Echinasteridae Echinodermata: Asteroidea ............ Echinodermata: Ophiuroidea .......... Elastic-sided gumleaf, or: the rubber cuticle and other studies of the Corymbo- sae, by S. G. M. Carr & D. J. Carr .... Erythraeidae Estuarine foraminiferal communities in Lake Illawarra, N.S.W., by I. Yassini & BuiGe JOMES Ignis eee ceases 297 205 206 207 207 208 208 209 210 210 101 327 159 297 27 382 374 375 369 407 159 267 101 193 229 PROC. LINN. SOC. N.S.W., 110 (3), (1987) 1988 Eucalyptus, host to Glycaspis I Seis Sh ONO oe i AE eee 11, 19, 25 Eucalyptus (Corymbosae), rubber cuticle .... 101 EUCaL) DiUSMOTCAd eS eee ae Ree 141 Farrant, P. A., & King, R. J., The Dictyo- tales (Algae: Phaeophyta) of New South Walesittogt stich ayer sce: seers usu 369 Hineandivecetatlonter sam ee err 146, 317 First report of Late Devonian trilobites from eastern Australia, by A. J. Wright 27 Fish, dietary study of golden-lined whiting 215 Fishilarnvac;warumeem scutes aris lo ci ceie nen 1 Floodplain waterbodies, crustacean ADO DAM AON scagnonaasososavedecne 297 Fluxes of inorganic nitrogen between sedi- ments and water in a coral reef lagoon, by R. Johnstone, K. Koop & A. W. D. WATT be erene etc eon Rt in eater 219 Foraminiferal communities, Lake Illa- Wel At geet cee sis yc ee ee oe ee eA 229 Glasby, P., Selkirk, P. M., Adamson, D., Downing, A. J., & Selkirk, D. R., Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the western Blue Mountains 141 Glycaspis/Eucalyptus associations ......... 19 Glycaspis (Synglycaspis) constricta sp. nov. ... 12 Glycasois (Synglycaspis) incomperta sp. nov. . . 25 Glycaspis (Synglycaspis) nancyana sp. nov. .. . 15 Glycaspis (Synglycaspis) rupicolae sp. nov. .. . 13 Golden-lined whiting ................. 215 Hemucytheridea hiltoni sp. nov. ........-.-. 171 Fiolocenewesetatlonuyrne eric tna 317 iomoeostrichuswolsentiy salmon ee ele ae 399 Homoeostrichus sinclairit ...............-. 400 Homoptera: Spondyliaspididae ....... 11, 19, 25 Hotchkiss, A. T., & Imahori, K., A new species of Nitella (Characeae) belonging to the pluricellulate species group in Australias ge stich) ins auoais arenes 175 Hotchkiss)3 AG ee é& "Imahori= Ke Additional observations on Nitella ver- ticillata (Characeae) from a new locality in Newssouth Walestiaee 2-922 187 Hunter Valley, crustacean zooplankton... 297 Hutchings, P. A., van der Velde, J. T., & Keable, S. J., Baseline survey of the benthic macrofauna of Twofold Bay, N.S.W., with a discussion of the marine species introduced into the bay ...... 339 Ichthyoplankton, Botany Bay........... 1 Imahori, K., see Hotchkiss, A. T. Inversa, group of Neurochaeta (Diptera) .... 49 Johnstone, R., Koop, K., & Larkum, A. W. D., Fluxes of inorganic nitrogen between sediments and water in a coral reehlagoonlss to nS ens eice ee eld. 219 Jones, B. G., see Yassini, I. PROC. LINN. SOC. N.S.W., 109 (3), (1987) 1988 Keable, S. J., see Hutchings, P. A. Kennedy, C. M. A., Pycnodithella harveyi, a new Australian species of the ‘Tri- denchthoniidae _(Pseudoscorpionida: /NGNG WONG EY) Su diee ds Colds Bald NMA 68 0 289 King, R. J., see Farrant, P. A. Kodela, P. G., & Dodson, J. R., A late Holocene vegetation and fire record from Ku-ring-gai Chase National Park, INW SOU WANES seeodccoeaccosoce 317 Koop, K.., see Johnstone, R. Ku-ring-gai Chase National Park, late Holocene vegetation ............... 317 Lacewing, a new species............... 407 Lake Illawarra, foraminiferal communities D229 Lake Munmorah, charophytes.......... 187 Larkum, A. W. D., see Johnstone, R. Late Holocene vegetation and fire record from Ku-ring-gai Chase National Park, New South Wales, by P. G. Kodela & JERS Dodson estos: cece 317 Lichen genus Cladonia section Cocciferae in Australia, by A. W. Archer.......... 205 Limnology ign as sce es va eee 297 Linnean Society of New South Wales. Record of annual general meeting 1986. Reports and balance SHEEES eee ee tee eee etches annexure 1 Bobophorayvaneoaias sss sae ee 393 Hobospirakbicuspidatas sy. 14) eee 381 McAlpine, D. K., Studies in upside-down flies (Diptera: Neurochaetidae). Part I. Systematics and phylogeny.......... 31 McAlpine, D. K., Studies in upside-down flies (Diptera: Neurochaetidae). Part II. Biology, adaptations, and specific mating mechanisms ............... 59 Macleay Memorial Lecture 1987 ........ 101 Magnifica, new group of Neurochaeta (Dip- tera ied ot Nea ao aes eee oer ae em 49 Melita plumulosaisps novi...) 1-00 1a 329 Mites, callidosomatine ................ 193 Moore, K. M., A new species of Glycaspis (Homoptera: Spondyliaspididae) and some new host records ............. 25 Moore, K. M., Associations of some Glycaspis species (Homoptera, Spondy- liaspididae) with their Eucalyptus species hosts¥sc8 SAS ea ee 19 Moore, K. M., see Campbell, K. G. Myrmeleontidae (Neuroptera) ......... 407 Nepean River, charophytes ............ 175 Neurochaeta, subgen. s. str. of Neurochaeta McAlpine e-em okie aan 44 Neurochaeta capilo sp. nov. ...........4... 46 Neurochaeta macalpinet ...............05. 52 Neurochaeta magnifica sp. nov. ............ 49 Neurochaeta parviceps sp. NOV. .........05. 52 Netrochactasabroshijten ect pea ee 52 INeuirochactidaemnse aero Neurocytta, subgen. nov. of Neurochaeta INICA pin ese ees tees ier ete eatce cease ase Neuroptera: Myrmeleontidae .......... Neurotexis, subgen. nov. of Neurochaeta INCA DINE eyes este e he cys eet acbnets New Australian larval callidosomatine mite (Acarina: Erythraeidae) parasitic on flies, with notes on subfamily and tribe classification, by R. V. Southcott ..... New genus and four new species in the family Echinasteridae (Echinoder- mata: Asteroidea), by F. W. E. Rowe & EapleewA be rtsOmese cer atee-cre eater rae New species of Melita (Crustacea: Amphi- poda: Melitidae) from northern New South Wales with a note on the genus Abludomelita Karaman, 1981, by W. TAENGNSO! ciorsg aos ens alaoeses o ein ener eae New species of Nitella (Characeae) belong- ing to the pluricellulate species group in Australia, by A. T. Hotchkiss & K. Imahori WNitellahverticillataeenpies eel sacle INotellarwoodttsps MOV) esa) oe ee Nitrogen flux between sediments and water Nothoasteia clausa sp. nov. ..........-.-.- Nothoasteia platycephala ................. Odontohenricia gen. nov. .............-.. Odontohenricia clarkae sp. nov. ........-.-. Odontohenricia endeavourt sp. nov. ......... Odontohenricia fishert sp. nov. ........-.-. Odontohenricia hayashii sp. nov. .........-. One Tree Island, nitrogen flux .......... Ophtopeza cylindrica............-+.4+0--- Ophiopeza spinosa Ophiopsammus aequalis Ophiopsammus anchista Ophiopsammus angusta sp. nov. ........... Ophiopsammusrassimilts sey vee oe Ophiopsammusiyoldi sa. 3. oe en Ophiuroidea Pachydictyon paniculatum ................ Padina australis Padina crassa RAGIN GM TAS CTs ari SRN sees Cea mee A GtnaNlenulsertete ines haciun took catane Parviceps, new group of Neurochaeta (Dip- tera) Pease, B. C., see Steffe, A. S. Rela gicye Osea eect mtv ancien s sess ts Phaeophyta (Algae), Dictyotales ........ Phytoglyphs of eucalypts .............. Platycopida Port Hacking, Recent ostracodes........ Portulacaceae, seed types in Calandrinia . . . Pseudoscorpionida: Arachnida ......... Pycnodithella harveyi, a new Australian species of the ‘Tridenchthoniidae 193 83 327 (Pseudoscorpionida: Arachnida), by (re Vitg Aue enn ec yan aera ener Rowe, F. W. E., & Albertson, E. L., A new genus and four new species in the family Echinasteridae (Echinoder- mMata-wASterOldea)) saya aes ereaeine Rowe, F. W. E., see Vail, L. L. Rubber cuticle of the Corymbosae......... Seed type and seed surface patterns in Calandrinia sens. lat. (Portulacaceae), by Syeda Saleha Tahir & R. C. Carolin Selkirk, D. R., see Glasby, P. Selkirk, P. M., see Glasby, P. Semicytherura illertt sp. nov. .....-....---. Sillago analis, dietary study ............. Smithers, C. N., A distinctive new species and new distribution records of Stilbop- teryginae (Insecta: Neuroptera, Myrmeleontidae) Southcott, R. V., A new Australian larval callidosomatine mite (Acarina: Erythraeidae) parasitic on flies, with notes on subfamily and tribe classifi- cation Spatoglossum macrodontum ..............- Spondylhiaspididae .................. 11 Status of the genera Ophiopeza and Ophio- psammus Echinodermata: Ophiuroidea) in Australian waters, with the descrip- tion of a new species, by L. L. Vail & F. WES ROWeseem acc atns posit en leun i ecninteg Steffe, A. S., & Pease, B. C., Diurnal survey of ichthyoplankton abundance, distri- bution and seasonality in Botany Bay, New South Wales Stulbopteryginae, distribution records .... Stilbopteryx mouldsorum sp. nov. .........- Studies in upside-down flies (Diptera: Neurochaetidae). Part I. Systematics and phylogeny, by D. K. McAlpine ... Studies in upside-down flies (Diptera: Neurochaetidae). Part II. Biology, adaptations, and_ specific mating mechanisms, by D. K. McAlpine ..... Study of the crustacean zooplankton of six floodplain waterbodies of the lower Hunter Valley, N.S.W., by B. V. Timms Stypopodium flabelliforme .............-.- Syeda Saleha Tahir & Carolin, R. C., Seed type and seed surface patterns in Calan- drinia sens. lat. (Portulacaceae) ...... WCOTTORAUSUTGIAS(CO eee Timms, B. V., A study of the crustacean ’ zooplankton of six floodplain water- bodies of the lower Hunter Valley, N.S.W. Tridenchthoniidae, anew species........ UmlobitesseateiDevonianweeeeeene ee Twofold Bay, benthic macrofauna ....... 289 83 101 307 172 215 407 193 386 5S), 267 297 401 307 404 297 289 27 339 PROC. LINN. SOC. N.S.W., 109 (3), (1987) 1988 Wipside-downbtlies ae eee: Vail, L. L., & Rowe, F. W. E., Status of the genera Ophiopeza and Ophiopsammus (Echinodermata: Ophiuroidea) in Aus- tralian waters, with the description of a NEWASPEClESHA EEE eC ee van der Velde, J. T., see Hutchings, P. A. Vegetation history, Ku-ring-gai Chase INationalwPark iin sbi il shire gett Warburton, K., see Brewer, D. T. Whiting, golden-lined (Szllago analis) ..... Wright, A. J., First report of Late Devonian trilobites from eastern Australia ..... Wright, A. J., see Yassini, I. 267 317 215 27 PROC. LINN. SOC. N.SW., 109 (3), (1987) 1988 Yassini, I., & Jones, B. G., Estuarine foraminiferal communities in Lake Weer INS obo oba eb eb doonacas Yassini, I., & Wright, A. J., Distribution and ecology of Recent ostracodes (Crustacea) from Port Hacking, New South Wales) se usanose Vee Zeidler, W., A new species of Melita (Crustacea: Amphipoda: Melitidae) from northern New South Wales with a note on the genus Abludomelita Kara- mani 1981) (ote a ah Ski ee ee Zonariaiangustata pee eee AQHA LGRHE oog060900000000000000n0¢ 229 159 327 394 395 397 4 f ADVIGEMLO AUTHORS The Linnean Society of New South Wales publishes in its Proceedings original papers and review articles dealing with biological and earth science. Papers of general significance are preferred but the Proceedings also provides a medium for the dissemination of useful works of more limited scope. Any work considered for publication in the Proceedings is subject to careful scrutiny by one or more external referees. Manuscripts will be received for assessment from non-members as well as members of the Society though non-members must communicate their works through a member. 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Like illustrations, they should be designed to fit a single page of the journal. fy 3 8 0 0 2 re Ee PROCEEDINGS of LINNEAN SOCIETY OF NEW SOUTH WALES VOLUME 110 Issued 13th January, 1989 CONTENTS: NUMBER 3 219 R. JOHNSTONE, K. KOOP and A W. D. LARKUM Fluxes of Inorganic Nitrogen between Sediments and Water in a Coral Reef Lagoon 229 |. YASSINI and B. G. JONES Estuarine Foraminiferal Communities in Lake Illawarra, N.S.W. 267 L. L. VAIL and F. W. E. ROWE Status of the Genera Ophiopeza and Ophiopsammus (Echinodermata: Ophiuroidea) in Australian Waters, with the Description of a new Species 289 C. M. A. KENNEDY Pycnodithella harveyi, a new Australian species of the Tridenchthoniidae (Pseudo- scorpionida: Arachnida) NUMBER 4 297 B. V. TIMMS A Study of the Crustacean Zooplankton of six Floodplain Waterbodies of the lower Hunter Valley, N.S.W. 307 SYEDA SALEHA TAHIR and R. C. CAROLIN Seed Type and Seed Surface Patterns in Calandrinia sens. lat. (Portulacaceae) 317 P. G. KODELA and J. R. DODSON A Late Holocene Vegetation and Fire Record from Ku-ring-gai Chase National Park, N.S.W. 327 W. ZEIDLER. A new Species of Melita (Crustacea: Amphipoda: Melitidae) from northern New South Wales with a Note on the Genus Ab/udomelita Karaman, 1981 339 P. A. HUTCHINGS, J. T. VAN DER VELDE, and S. J. KEABLE Baseline Survey of the Benthic Macrofauna of Twofold Bay, N.S.W., ‘with a Discussion of the Marine Species introduced into the Bay 369 P A. FARRANT and R. J. KING The Dictyotales (Algae: Phaeophyta) of New South Wales 407 C.N. SMITHERS A distinctive new Species and new Distribution Records of Stilbopteryginae (Insecta: Neuroptera: Myrmeleontidae) 411 INDEX Printed by Southwood Press Pty Limited, 80-92 Chapel Street, Marrickville 2204 I » i MAR ee ete f : fe : 4 eK ait ” ‘i te ny ait ' y 7 ; i ; i t ( , f ; ) : 1 i) it, ; \ hy : , } i i aa k , . t fi i hy f it + ft x j Pr ’ j i A ‘ i j f ve ; f ay 5; ney ied - t i bad y tat SFp eat Ta nar wy a, rs ave ven cre ts Cres Se ee ee > LS rate hy PY $F fret ns ; ! » ait. shiek b oe tend ate 4 : : Rin4) A és rs bein ees i te fgievid Shee wn te tind LSAT DP 0E 7 ed f LSiendety wed \ eit ose (eh arbre BE MEG OPE 4 sorn eth ot Parner ar) FF AE UREN SVL A Sh M4 « 2 oe ver oY Cet MN reer Oat Rie ed el kal Blt ed Bad lick 4 ; HP vee hadehe petieg A Rone Hl eG uty tee f eee ed Lge Btowat et ‘ f ‘ ‘ ou errr Stern “ * Le the d sche : teh Be ‘ 2 : pare hots eerie r sl ee Ke aw foe < { Pate : A HRLER TAR CH ES aber 4 i d ‘ poh dt age ADE Sh etn : Ws vs estep Samleua toh Bats f bedi ye ib ak cG ree TO CIM B® 44,02 ap Beh E A Oy Serer ner io tat! © a ey a 4 Hy 4 Rises ies U aes “fh ERTS, ste a mde noni oY “i d x ere ie et ene aD etteh wot eet et HiRes eyed eee & ZB) } Bice hot Oe eS ; EBs Fo Svat et hd Chey pueda) Saitaty (Oh oh gfe Ot Co nb eR at 5 LE eb Myonsthoy (oe it eat shane pie hdl meh he fox Boe tek ety net teatict es aves rbo® ORE BR EME TL x Vaget storms He arn ra gasetat embers : Panett S Sp ieebat eek eich @ Wied ohois ss OPER E LAT eet Hote eh Disa one seme a R Leper it : wage Prat : 4 Gin Coe an a th 4 ehonsb doeshes rie } a ofiee Se RSD aE A Pitre BO Tesrges soG ee Tarr “, at, enh ten fener hotest * Cet ne rae srr see eee Lata ae: tak ceesus es sie a nak eh tt : smth Fa AEE SEY A4 YOO Sh ata ee A pesar gp Aloe yuan eh REE AE ar ee 2 ae fee te Eee pte eet ee hots Par Se a Sere ee ee é ” Tate eee ee ne d are ee pone inerteteneses € oie Oe ee rn, Per Ca gar fererrs ey wrap fares tr are mene Or be eases Be eese greene? 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