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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 Microbiology 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 $53.00 The current rate of subscription to the Proceedings for non- members is set at $80.00 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 Secratary. OFFICERS AND COUNCIL 2000-2001 President: J.P. BARKAS Vice-presidents: R.J. KING, K.L. WILSON, A. RITCHIE, R.A.L. OSBORNE Honorary Treasurer: 1.G. PERCIVAL Secretary: C.G. FORD Council: A.E.J. ANDREWS, M.L. AUGEE, J. BARKAS, M.R. GRAY, G.J. HARDEN, M.S. MOULDS, D.R. MURRAY, P.J. MYERSCOUGH, R.A.L. OSBORNE, I.G. PERCIVAL, A. RITCHIE, S. ROLE, K.L. WILSON Honorary Editor: M.L. AUGEE Linnean Macleay Lecturer in Microbiology: P.R. REEVES Auditors: Phil Williams Carbonara The postal address of the Society is P.O. Box 137, Matraville, N.S.W. 2036, Australia. Telephone and Fax +61 2 9662 6196. E-mail: linnsoc @acay.com.au Home page: http://www.acay.com.au/~linnsoc/welcome.html ©) Linnean Society of New South Wales Cover motif: An onychophoran from eastern Australia. Eight new onychophorans are described in this volume. PROCEEDINGS of the LINNEAN SOCIETY NEW SOUTH WALES For information about the Linnean Society of New South Wales, its publications and activities, see the Society’s homepage http://www.acay.com.au/~linnsoc/welcome.html VOLUME 122 December 2000 Proc. Linn. Soc. N.s.w., 122. 2000 fy : : p x 2 if, + re Be ‘ ~T Eight New Planipapillus (Onychophora: Peripatopsidae) from Southeastern Australia AMANDA REID 140 Napoleon Street, Eltham, Victoria 3095 E. mail: mandyr@connexus.net.au Reid, A. (2000). Eight new Planipapillus (Onychophora: Peripatopsidae) from southeastern Australia. Proceedings of the Linnean Society of New South Wales 122, 1-32. Eight new Planipapillus Reid, 1996 (Onychophora) are described from southeastern Australia: P. annae, sp. nov.; P. berti, sp. nov.; P. biacinoides, sp. nov.; P. cyclus, sp. nov.; P. gracilis, sp. nov.; P. impacris, sp. nov.; P. tectus, sp. nov. and P. vittatus, sp. nov. The generic diagnosis is emended to add some characters that were not included in the original diagnosis. Planipapillus are widespread throughout the southern highlands of Australia and the adjacent escarpment and it is likely that many more, particularly cryptic species, await discovery. Manuscript received 14 September 1999, accepted for publication 19 April 2000. KEY WORDS: Peripatopsidae, Planipapillus, southeastern Australia, taxonomy. INTRODUCTION Australia has more described species of Onychophora (Peripatopsidae) than any other continent. It remains to be seen whether this is a reflection of a focused research effort in this country in recent years, or a true representation of the biogeography of the group. It appears that New Zealand may also have a rich onychophoran fauna (Tait and Briscoe 1995; Gleeson 1996; Trewick 1998). The Australian fauna was recently reviewed by Reid (1996). Fifty six nominal species were recognised in that paper, though many were identified as belonging to probable cryptic species complexes. This was recently confirmed for one putative species, Euperipatoides rowelli Reid, 1996 using microsatellite markers (Sunnucks and Wilson 1999) and has been suggested for other species on the basis of numerous allozyme studies (Briscoe and Tait 1993). Thus, the true number is probably far greater than the tally of nominal species due to high levels of local endemism in the family, morphological crypsis, and the vast extent of hitherto unexplored potential onychophoran habitat in this continent. New members of a single genus, Planipapillus Reid, 1996 are described here. This genus is restricted to southeastern Australia and its members are found from the southern highlands, at altitudes of up to 1,737 m within the treeline leading up to Mt Kosciuszko (2,228 m), to the eucalypt forests of the adjacent escarpment in the southeast corner of the country. Like other onychophorans, they are found in humid micro-habitats; occurring primarily in and under decomposing logs and leaf litter in a range of forest types, including remnant rainforest pockets (such as the Tarra-Bulga National Park in South Gippsland) as well as open dry sclerophyll forests. Onychophorans are commonly thought to occur in lush, moist forest habitats but in many cases the macro-habitats where Planipapillus spp. are found are relatively dry and may be far from pristine (Fig. 6). Proc. Linn. Soc. n.s.w., 122. 2000 bo NEW PLANIPAPILLUS (ONYCHOPHORA) Planipapillus was so-named to highlight a very characteristic trait of its members: a patch of reduced papillae on the heads of males. Modifications of the heads of male Peripatopsidae are postulated to be secondary sexual characters with a possible role in the transfer of spermatophores to females. Spermatophores have been found associated with these structures in some species (Tait and Briscoe 1990) and mating has been observed on two occasions: in an undescribed species (Meredith 1995, Tait pers., comm.) and in Planipapillus annae, sp. nov. described here. The author’s (1996) revision was based primarily on museum specimens and extensive collecting throughout New South Wales. Specimens from southern Australia were poorly represented in these collections. The present study is part of a wider project to redress this problem through the collection and identification of onychophorans from southeastern mainland Australia. Only Planipapillus species are included in this paper, though members of this genus were often collected with undescribed Ooperipatus Dendy, 1900. Members of the latter genus, will be described elsewhere. Four species of Planipapillus were described previously: P. biacinaces, Reid, 1996; P. bulgensis; Reid, 1996; P. mundus Reid, 1996 and P. taylori Reid, 1996. Eight new species are described below. MATERIALS AND METHODS Specimen Collection and Preservation This study is based on the examination of preserved specimens, most of which were hand collected from within and under decomposing logs. Specimens were preserved partially following the method of Reid (1996). Animals were anaesthetised by exposure to ethyl acetate vapour for 10 min, dipped in 70% ethanol to render the cuticle less hydrophobic, and fixed in 4% formalin for 2—3 days, then stored in 70% ethanol. Animals preserved in this way are distended, enabling characters to be examined more easily than is possible in contracted specimens. Tissue Preparation for Transmitted Light Microscopy Cuticular tissue was cleared in a small volume (approximately 1 ml) of saturated potassium hydroxide (KOH/H,O) solution on a hotplate set at approximately 50°C. Following clearing (approximately 1 hr; tissue appears translucent), a drop of 1% aqueous aniline blue was added with sufficient lactic acid (approximately 2 drops) to neutralise the solution, rendering the aniline blue the correct colour for staining. [In extreme alkaline conditions (KOH/H,O solution) aniline blue appears red; neutralising the solution, or making it slightly acidic, restores the blue colour of the stain.] Tissue pieces were stained for 15 mins, rinsed in water and mounted in glycerol jelly. The stained and mounted tissue was examined using a compound microscope and drawings made using a camera lucida. This method differs slightly from that of Reid (1996). Washing tissue pieces after clearing, as detailed in Reid (1996), has proved very difficult (cleared tissue pieces are difficult to see, and consequently often lost during washing) and time consuming, so this step has been eliminated from the method. For males and females of each species, the following tissue samples were prepared as above: dorsal integument, nephridiopores, crural papillae (where present) from oncopods 3, 7 and 12, and anterior accessory gland papillae and posterior accessory gland foramen. Unfortunately, no diagnostic differences were found in these characters among the eight species described (and the four Planipapillus described in Reid, 1996), so few are illustrated. Terminology Terminology for all characters follows Reid (1996). Head width is used as an indicator of size as this measure is less prone to variation due to the degree of distension Proc. Linn. Soc. n.s.w., 122. 2000 A. REID 3 of the body than are other size indicators, such as total length. Where measurements and counts are given, these refer only to type specimens. Measurement values are expressed as minimum-mean-maximum. Abbreviations EDI eye diameter index, expressed as a proportion of head width HWE width of head measured dorsally between the midpoint of each eye MV Museum Victoria Taxonomy Morphological variation within and among populations was assessed to identify species. A phylogenetic analysis based on morphological characters was attempted, but no clades within Planipapillus were clearly resolved. As many characters that differ among species relate to the male modified head papillae, it is difficult to be satisfied that assumptions of character independence can be met when conducting such an analysis. The generic description given below comprises characters present in all Planipapillus species. Only characters that differ among species are given in the species descriptions. Species descriptions were generated using DELTA (Description Language for Taxonomy) software (Dallwitz 1980; Partridge et al. 1993; Dallwitz et al. 1993). TAXONOMY Genus Planipapillus Reid Planipapillus Reid, 1996: 851-852, fig. 30. Type species: Planipapillus taylori Reid, 1996: 853-856, Figs 71, 119, 120. Type: holotype o, (AM KS40020). Diagnosis (emended from Reid, 1996) Colour pattern comprises: longitudinal light-coloured band along dorsal midline and short, dark, transverse bars or blotches along midline dorsal to oncopods; light dorsolateral transverse patches in line with oncopods and light patches laterally between oncopods (components are variably present within and among species). Males with an ovoid patch of reduced papillae posterior to eyes. Vas deferens continues directly (without looping posteriorly) from vasa efferentia to gonopore. Females with, or without crural papillae. Oviparous. Generic description Colour pattern Body pigmented. Pigment not soluble in alcohol. Primary papillae light basally, dark tipped; longitudinal light-coloured band along dorsal midline and short, dark, transverse bars or blotches along midline dorsal to oncopods; light dorsolateral transverse patches in line with oncopods and light patches laterally between oncopods (components are variably present within and among species). Ventral organs whitish. Oncopods colour similar to, or slightly paler than body. Antennal rings Approximately 30 antennal rings in adults and juveniles; wide and narrower antennal rings alternate; each with single row of bristles; proximal antennal rings expanded ventrally to form sensory pads. Eyes Present. Head (males) Modified papillae (i.e. different from remaining dorsal papillae) present; an ovoid patch of reduced papillae posterior to eyes; papillae adjacent to patch usually enlarged. No eversible head structure, furrow between antennae, or modified papillae anterior to eyes. Proc. Linn. Soc. N.s.w., 122. 2000 aS NEW PLANIPAPILLUS (ONYCHOPHORA) Jaws Inner jaw with 4—6 (usually 5) denticles; diastema absent. Outer jaw with accessory tooth. Integument Dorsum with 12 complete plicae between oncopods; wide and narrow plical folds alternate. Papillae not uniform in size, alternate plicae with some slightly larger, usually primary papillae. Papillae arrangement: primary papilla with short, narrow bristle between pair of larger primary papillae with longer, more robust bristles and smaller secondary papillae between primary papillae; conical apical piece absent; papillar scales ribbed in both sexes, remaining integument with small scales. Lateral primary papillae slightly enlarged or elongate, with more prominent pair between oncopods in line with junction of oncopods and body. Size of papillae posterior to gonopore similar to rest of ventrum, approximately same size dorsally and ventrally, with only those surrounding anal opening enlarged. Oncopods Number of pairs of oncopods invariant intraspecifically; 15 pairs in both sexes. Last pair of oncopods fully developed in both sexes; orientation as for remaining oncopods. First pair of oncopod feet not enlarged, similar in size to remaining feet. Basal foot papillae absent. Distal foot papillae present, one anterior, one median, one posterior; each papilla with single sensory bristle. Oncopods with three complete spinous pads; fourth broken spinous pad present; spinous pads well-developed on all oncopods. Nephridiopores at center of third spinous pad on fourth and fifth oncopod pair; nephridiopore openings crescent-shaped, surrounded by smooth lip. Male reproductive tract Gonopore (both sexes) between last pair of oncopods. Male genital pad low, rounded, not protuberant or penis-like; composed of large papillae with ribbed scales; papillae sometimes fused surrounding gonopore; gonopore shape cruciform (with arms equidistant), arms extending close to rim of genital pad. Vasa efferentia with thin flexible walls; proximal vasa efferentia broad; vas deferens not thick walled, opaque, not shiny. Spermatophore pouch present. Male glands and gland papillae Crural glands and crural papillae present. Crural papillae protrude between plicae 4_5 (counting from third spinous pad); with finely ribbed scales basally, distally scales broad, with distinct ribs; open via short slit; smooth rim surrounding distal foramen ovoid or lip-shaped, not extending to papilla margin. Some crural glands extend from oncopods into lateral haemocoel, while others are confined within oncopods. Coxal organs absent. Anterior accessory gland papillae present, or absent; if present, open on genital segment at base of last pair of oncopods via longitudinal slit. Anterior accessory glands present, or absent. Posterior accessory glands present; open directly to exterior on anal segment approximately midway between genital and anal openings; gland foramen separate, close together; glands broad and saccate. Female reproductive tract Females with ovipositor; oviparous; gonopore shape longitudinal slit. Ovarian tubes separate, suspended along entire length to pericardial floor; with thin walls; oviducts unite close to ovary. Ova follicular; large, yolky. Spermathecae present, well-developed. Receptaculum ovorum absent. Additional pouches present. Remarks The generic diagnosis has been emended to include characters relating to the colour pattern, and the presence of crural papillae in females. Features of the colour pattern are very distinctive in members of this genus. These were not included in the original diagnosis. However, with the description of more Planipapillus species, it has become apparent that components of the pattern are present in all members of the genus known thus far and are useful traits to recognise, and help define the genus. Proc. Linn. Soc. n.s.w., 122. 2000 A. REID >) The possession of crural papillae in females, although not present in all species, is a trait that occurs in only two other onychophoran genera, Peripatopsis Pocock, 1894 and TJasmanipatus Ruhberg et al., 1991, and is, therefore, useful to include in the generic diagnosis for Planipapillus. The posterior section of the male reproductive tract, showing the vas deferens and part of the vasa efferentia, is illustrated for each species (Fig. 4). [Complete tracts are shown for P. berti, sp. nov. (Fig. 9a), and P. impacris, sp. nov. (Fig. 16).] While some slight differences are apparent, the overall structure is very similar in each species. When the tracts are swollen with sperm, the vasa efferentia are usually separate, but in other specimens the vasa efferentia lie parallel for part of their length (compare Figs 4h and 41, and Figs 9a and 16). In all species, the vas deferens continues directly from the joined vasa efferentia to the gonopore. This trait is characteristic for members of this genus. In most other Peripatopsidae, the vas deferens continues anteriorly for a short distance before looping, hairpin-like, posteriorly toward the gonopore following the junction of the vasa efferentia. Planipapillus annae, sp. nov. (Figures la; 1b; 2a; 3; 4a; 5a; 5b; 7. Table 1) Material examined Holotype: ©% Victoria, 5.9 km NW Bonang, beside Deddick R. Rd (between Bonang and Tubbut), 37°11’S 148°41’E, 740 m, 14 Jun 1999, coll. A. Reid and A. Skates (MV K7281). Paratypes: Victoria, 50} 59, data as for holotype (MV K7282). Diagnosis Body with mid-dorsal dark stripe; without median longitudinal light-coloured band; antennal rings not banded. Ovoid patch of reduced papillae on heads of males without sclerotised spikes; 21-24 rows of plicae comprise patch; 2—3 rows papillae lateral to patch elongate, all similar length, each with single sensory bristle. Anterior accessory glands and gland papillae absent. Posterior accessory glands straight or folded distally, short hook. Description Measurements HWE males 0.87—0.91—1.00 mm (n=5, Holotype 0.90 mm HWE); HWE females 0.87—0.97—1.00 mm (n=5). Colour pattern Ground colour greyish-blue, or olive green. Mid-dorsal dark stripe present; short, dark, transverse bars or blotches along midline dorsal to oncopods (indistinct in dark specimens); evenly scattered tan or tan-based papillae (Fig. 1a); laterally with longitudinal light band dorsal to oncopods, or with light patches between oncopods (indistinct). Oncopods with cream patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present, very pale. Spinous pads greyish- blue. Integument between genital and anal openings darker than rest of ventrum. Antennal rings not banded, ground colour. Antennal rings Distal 8-9 antennal rings with sensory bulbs; sensory pads with two rows of sensilla. Eyes EDI males 0.06—0.06—0.07; EDI females 0.06—0.06—0.07. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in longitudinal ovoid patch posterior to eyes (Figs 1b, 2a and 3). Patch without sclerotised spikes at medio-posterior margin. Ovoid patch comprising 21—24 Proc. Linn. Soc. N.s.w., 122. 2000 Table 1. Planipapillus spp. distinguishing features. Symbols and abbreviations : A = Absent; B = banded; BL = mid-dorsal blotches dorsal to oncopods; BO = light band laterally above oncopods; C = cylindrical papillae; DS = mid-dorsal dark stripe; HP = male patch of modified head papillae; LB = longitudinal tan or light brown band along dorsal midline; NB = not banded; P = Present; PO = light patches laterally between oncopods; S = semicircular papillae; TP = pale dorso-lateral transverse patches; * not all specimens examined show all traits; # numbers refer to oncopod numbers. le crural nterior Posterior a Papillae emale rings rows of papillae rows / bristles within patch papillae/ crural _ accessory gland accessory accessory crural plicae shape (enlarged glands extending papillae glands glands papillae # papillae) into lateral haemocoel # NEW PLANIPAPILLUS (ONYCHOPHORA) P. annae, sp. nov. 2-3 laterally; T 2-3 and 6-14/ folded, short posteriorly / C ridge I-14 hook or straight, blunt P. berti, sp. nov. BL, LB, PO B A 18-22 1-2 laterally, up to 3 short, smooth 2-3 and 6-14/ A (usually) A straight, A inner row longest ridge 1I-14 blunt IC P. biacinaces Reid, BL, LB, PO, TP B 2 short 10-11 2-3 laterally / C 1 low, smooth, 2-3 and 6-14/ P greatly reduced folded, short 2-3(?) and 1996 conical 11-14 hook 6-14 P. biacinoides, sp. BL, LB, PO, TP B, NB 2 short 6-7 2-3 laterally / C 1 low, smooth, 2-3 and 6-14/ P greatly reduced folded, short P nov. conical 11-14 hook P. bulgensis, Reid, BL, LB, PO B A 16 2-3 laterally / C 1 short, smooth 2-3 and 6-14 A A straight, A 1996 ridge 11-14 blunt P. cyclus, sp. nov. BL, BO, DS, LB B, NB 4 subequal 6-7 1, entire margin / up to 4 greatly 2-3 and 6-14/ P short folded, long 2-3 and 6-14 S reduced, 6-14 hook smooth P. gracilis, sp. nov. BL, PO B, NB A 19-20 1-2 laterally, up to 3 short, smooth 2-3 and 6-14/ A (usually), or A straight, A inner row longest; ridge I-14 greatly reduced blunt, or 1 posteriorly / C pointed P. impacris, sp. nov. BL, DS; LB, PO, B, NB 4 subequal 6-7 1 entire margin + up to 7 greatly 6-14/ P greatly reduced folded, long 6-14 TP 1 dorsolateral and reduced, 9-14 hook (LB and TP juvenile 1 ventrolateral smooth only) pair in 2nd row / ; S P. mundus Reid DS, LB, PO B A 19-24 2-3 laterally / C 1 short, smooth 1 or 2-3 and A A Straight, A 1996 ridge 6-14/ 11-14 blunt P. taylori Reid, BL, DS, LB, PO, B 4 equal 10-11 2-3 rows entire up to 15 short, smooth 6-14/ P greatly reduced folded, long 6-14 1996 TP margin / C ridge 6-14 hook P. tectus, sp. nov. BL, BO, LB, PO, B A 15-17 1 laterally (slight) 1 semicircular 2-3 and 6-14/ P greatly reduced folded, long 2-3 and 6-14 TP /C with scales 11-14 hook or anteriorly; straight, short, smooth blunt ridge posteriorly P. vittatus, sp. nov. BL (trace), BO, LB, NB A 5-6 not enlarged up to 2 short, smooth 2-3 and 6-14/ P greatly reduced folded, long 6-14 PO, TP (TP juvenile ridge I-14 hook only) Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 7 Figure 1. Planipapillus annae, sp. nov.: (a) body, holotype male 0.90 mm HWE, scale bar 0.30 mm; (b) head, holotype male, 0.90 mm HWE, scale bar 0.30 mm. Planipapillus berti, sp. nov.: (c) body, paratype male, 0.85 mm HWE, scale bar 0.20 mm; (d) head, paratype male, 0.87 mm HWE, scale bar 0.20 mm. rows plicae; papillae forming patch triangular, ridge-like; papillar scales fused, papillae smooth, each with a sensory bristle. Papillae laterally and posteriorly adjacent to patch cylindrical, enlarged; 2—3 rows of enlarged papillae laterally, enlarged papillae all similar length, innermost row only slightly longer than rest; four enlarged papillae in single transverse row posterior to patch; each enlarged papilla with single bristle (Figs 1b and 3). Proc. Linn. Soc. n.s.w., 122. 2000 8 NEW PLANIPAPILLUS (ONYCHOPHORA) Head (females) Females with no modification of head papillae. Dorsal integument Males with 12—12—13, females with 12—13—14 papillae counted from mid- dorsal line to junction of oncopod 10. Primary papillae cylindrical. Male reproductive tract Male genital pad cylindrical, protuberant. Proximal vasa efferentia separate, do not lie parallel before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Fig. 4a). f g h Figure 2. Planipapillus, spp. nov. diagrams of male heads to show relative positions of patches of modified papillae (shaded): (a) P. annae, sp. nov.; (b) P. berti, sp. nov.; (c) P. biacinoides, sp. nov.; (d) P. cyclus, sp. nov.; (e) P. gracilis, sp. nov.; (f) P. impacris, sp. nov.; (g) P. tectus, sp. nov.; (h) P. vittatus, sp. nov. Proc. Linn. Soc. n.s.w., 122. 2000 A. REID 9 Figure 3. Planipapillus annae, sp. nov., modified region of head, holotype male, 0.90 mm HWE, scale bar 0.30 mm. Male glands and gland papillae Crural papillae on ventral side of oncopods 2-3 and 6-14. Papillae shape differs among oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section (oncopods 2-3) or semicircular or cylindrical, tapered slightly distally, not divided into distinct basal and distal regions (oncopods 6— 14); papillae oncopods 6-10 narrow, low, cylindrical. Crural glands extend into lateral haemocoel from oncopods 11-14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae absent. Anterior accessory glands absent. Posterior accessory glands straight, bulbous, blunt distally (Fig. 5a), or folded distally, short hook tapered only slightly to blunt tip (Fig. 5b). Female crural papillae Absent. Remarks Planipapillus annae, sp. nov. differs from most other species that do not have spikes on the posterior margin of the head patch in lacking the pale mid-dorsal longitudinal light-coloured band that is very distinctive in many members of this genus. While some P. mundus and P. gracilis, sp. nov. specimens also lack this band, P. mundus does not have dark mid-dorsal blotches dorsal to each oncopod, and has banded antennae, unlike P. annae, sp. nov. Planipapillus gracilis, sp. nov. differs in having up to three sensory bristles on the enlarged papillae lateral to the male head patch. The papillae on each side of the head patch are not as elongate in P. annae, sp. nov. as they are in P. berti, sp. nov. and P. gracilis, sp. nov., and are of similar length, while those of the latter two species vary in length. Additional characters that distinguish this species from other members of the genus are shown in Figs 2, 5 and Table 1. Proc. Linn. Soc. n.s.w., 122. 2000 10 NEW PLANIPAPILLUS (ONYCHOPHORA) Ep of ‘Bo Figure 4. Planipapillus, spp. nov. posterior section of male sua lal tracts showing vas deferens and part of vasa efferentia for comparison: (a) P. annae, sp. nov., vd, vas deferens, ve, vasa efferentia, n=6, scale bar 0.50 mm; (b) P. berti, sp. nov., n=12, scale bar 0.50 mm; (c) P. biacinoides, sp. nov., n=2 (in addition, one specimen with tracts similar in appearance to P. impacris, sp. nov., Fig. 4f), scale bar 0.30 mm; (d) P. cyclus, sp. nov., n=3, scale bar 0.30 mm; (e) P. gracilis, sp. nov., n=4, scale bar 0.30 mm; (f) P. impacris, sp. nov., n=2 (in addition one specimen with tracts similar in appearance to P. tectus, sp. nov., Fig. 4g), scale bar 0.50 mm; (g) P. tectus, sp. noy., n=4, scale bar 0.50 mm; (h) P. vittatus, sp. nov., n=1, scale bar 0.50 mm; (i) P. vittatus, sp. nov, n=2, scale bar 0.30 mm. n, number of male reproductive tracts examined exhibiting the illustrated shape. When specimens were being prepared for preservation following collection, two specimens were observed presumably mating. The head of a male specimen was attached to the genital opening of a female specimen and both walked around in this position for some time. The tip of the ovipositor of the female was positioned on the patch of reduced papillae on the male’s head and the elongate papillae on each side of the patch clasped the ovipositor. The female was not subsequently checked for the presence of sperm in the genital tract to determine whether insemination had taken place. “Head-to-tail’ mating has been observed in only one other species of onychophoran to date (Meredith 1995 and Tait pers. comm.). Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 11 Habitat In and under logs in dry sclerophyll woodland. Specimens were collected beside road in cleared farming area (Fig. 6). Live, hand collected specimens were usually coiled in spiral when first exposed. Distribution This species is known only from the type locality (Fig. 7). yy 4 BY YS ye yy Figure 5. Planipapillus, spp. nov. posterior accessory glands: (a) P. annae, sp. nov., n=5, scale bar 0.30 mm; (b) P. annae, sp. nov., n=7, scale bar 0.30 mm; (c) P. berti, sp. nov., n=24, scale bar 0.20 mm; (d) P. biacinoides, sp. nov., n=6, scale bar 0.30 mm; (e) P. cyclus, sp. nov., n=6, scale bar 0.40 mm; (f) P. gracilis, sp. nov., n=7, scale bar 0.20 mm; (g) P. gracilis, sp. nov., n=1, scale bar 0.20 mm; (h) P. impacris, sp. nov., n=6, scale bar 0.50 mm; (i) P. tectus, sp. nov., n=6, scale bar 0.30 mm; (j) P. tectus, sp. nov., n=1, scale bar 0.30 mm; (k) P. tectus, sp. nov., n=1, scale bar 0.30 mm; (1) P. vittatus, sp. nov., n=5, scale bar 0.20 mm. n, number of glands examined exhibiting the illustrated shape. (Where two glands are shown for a species, individuals may have one of each type.) Proc. Linn. Soc. N.S.W., 122. 2000 12 NEW PLANIPAPILLUS (ONYCHOPHORA) a ee. fi id Figure 6. Type locality of Planipapillus annae, sp. nov., beside Gelantipy Rd. between Bonang and Tubbut, Victoria. Anne Skates (after whom the species is named) is searching fallen timber for specimens. Etymology This species is named in honour of Anne Skates, who found the first specimen and has accompanied me on many fieldtrips in search of Onychophora. Bairnsdale « Figure 7. Distributions of Planipapillus spp.: (1) P. annae, sp. nov.; (2) P. berti, sp. nov.; (3) P. biacinaces Reid, 1996; (4) P. biacinoides, sp. novy.; (5) P. bulgensis Reid, 1996; (6) P. cyclus, sp. nov.; (7) P. gracilis, sp. nov.; (8) P. impacris, sp. nov.; (9) P. mundus, Reid, 1996; (10) P. taylori Reid, 1996; (11) P. tectus, sp. nov.; (12) P. vittatus, sp. nov. 1,000 m contour shown. Note: Specimens of P. biacinaces have now been found at Falls Creek. They were previously only known from the type locality, Howman Gap, Victoria. Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 13 Planipapillus berti, sp. nov. (Figures lc; 1d; 2b; 4b; 5c; 7; 8; 9. Table 1) Material examined Holotype: C Victoria, Granite Flat, 9 km S of Mitta Mitta, beside Omeo Hwy, N of intersection of Omeo Hwy and Walsh’s Rd, 36°35’S 147°27’E, 350 m, 9 Mar 1999, coll. A. Reid and R. Roberts (MV K7283). Paratypes: Victoria, 12c) 39, data as for holotype (MV K7284). Diagnosis Body with longitudinal light-coloured band along dorsal midline and dark bands or blotches dorsal to oncopods; antennae banded. Ovoid patch of reduced papillae on the heads of males without sclerotised spikes; 18—22 rows of plicae comprise patch; 1—2 rows of papillae lateral to patch elongate, finger-like, each with up to three sensory bristles. Anterior accessory glands absent and gland papillae usually absent. Posterior accessory glands straight, blunt. Description Measurements HWE males 0.80—0.87—0.97 mm (n=13, Holotype 0.97 mm HWE); HWE females 0.87—0.88—0.90 mm (n=3). Figure 8. Planipapillus berti, sp. nov., modified region of head, holotype male, 0.97 mm HWE, scale bar 0.20 mm. Colour pattern Ground colour greyish-blue (few specimens brownish). Mid-dorsal dark stripe absent; longitudinal light-coloured band along dorsal midline and short, dark, transverse bars or blotches along midline dorsal to oncopods [band light ground colour, or, in two of 16 specimens, tan (Fig. 1c)]; laterally with distinctive cream patches between oncopods. Oncopods with light patches at junction with feet. Papillae around anal opening pigmented as for rest of body, or sometimes pale yellow. Ventral pigment present, very pale ground colour. Spinous pads pale yellow, or greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum. Proc. Linn. Soc. N.s.w., 122. 2000 14 NEW PLANIPAPILLUS (ONYCHOPHORA) Antennal rings banded, tan or with tan mottle dorsally; dorsal banding on alternate rings distal to, and including ring five (very pale, mottling on basal third of antennae only). Antennal rings Distal 8—9 antennal rings with sensory bulbs; sensory pads with two rows of sensilla. Eyes EDI males 0.06—0.07—0.10; EDI females 0.07—0.08—0.08. Head (males) Males with modified papillae on head (1.e. differ from remaining dorsal papillae). Papillae reduced in longitudinal ovoid patch posterior to eyes (Fig. 2b). Patch without sclerotised spikes at medio-posterior margin. Ovoid patch comprising 18—22 rows plicae; papillae forming patch triangular, ridge-like (with 8—10 reduced papillae on each plica); papillar scales fused, papillae smooth, each with a sensory bristle. Papillae laterally adjacent to patch cylindrical, finger-like, enlarged; 1—2 rows of enlarged papillae, inner row longest with 7—10 papillae; long and shorter papillae alternate in length from anterior to posterior; each enlarged papilla with up to three bristles (usually 1—2) (Figs 1d and 8). Head (females) Females with no modification of head papillae. Dorsal integument Males with 12—12—14, females with 11—12—14 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae cylindrical. Figure 9. Planipapillus berti, sp. nov.: (a) male reproductive tract and associated glands, paratype, 0.87 mm HWE, scale bar 0.50 mm; (b) distal tip of crural papilla oncopod 3, paratype male, 0.85 mm HWE, scale bar 0.03 mm; (c) crural papilla oncopod 3, side view, paratype male, 0.85 mm HWE, scale bar 0.02 mm; (d) crural papilla oncopod 12, paratype male 0.85 mm HWE, scale bar 0.05 mm. cg, crural gland; pa, posterior accessory gland; sv, seminal vesicle; t, testis; vd, vas deferens; ve, vas efferens. Proc. LINN. Soc. n.s.w., 122. 2000 A. REID 15 Male reproductive tract Proximal vasa efferentia separate, do not lie parallel before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Figs 4b and 9a). Male glands and gland papillae Crural papillae on ventral side of oncopods 2-3 and 6—14. Papillae shape differs among oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section [oncopods 2-3 (Figs 9b and 9c)], or semicircular or cylindrical, tapered slightly distally, not divided into distinct basal and distal regions [oncopods 6—14 (Fig. 9d)]. Crural papillae oncopods 2-3 large; crural papillae small on oncopods 6-9, 10 or 11 (variable); remaining crural papillae large. Crural glands extend into lateral haemocoel from oncopods 11-14; straight, short, not folded (Fig. 9a); remaining glands confined within oncopods. Anterior accessory gland papillae usually absent. Anterior accessory glands absent. Posterior accessory glands straight, tapered to blunt point distally (Figs 5c and 9a). Female crural papillae Absent. Remarks The papillae lateral to the head patch in male Planipapillus berti, sp. nov. are much longer than those in all other species, with the exception of P. gracilis, sp. nov. Planipapillus berti, sp. nov. differs from P. gracilis, sp. nov. in having a longitudinal light-coloured band along the dorsal midline. Additional characters that distinguish P. berti, sp. nov. from other Planipapillus are given in Figs 2, 5 and Table 1. Usually a greater number of female than male Planipapillus are found at a collection site, suggesting a sex ratio biased towards females in Planipapillus populations. At Granite Flat, the converse was true; 14 of a total of 20 specimens collected were male. Whether this ratio is a true representation for the entire population is unknown. A single male specimen had an anterior accessory gland papilla (but without a corresponding anterior accessory gland) on one of the last oncopods. Planipapillus berti, sp. nov. was found with two male Ooperipatus sp. Habitat In and under logs in dry sclerophyll woodland. Live, hand collected specimens were usually coiled in spiral when first exposed. Distribution The species is known only from the type locality (Fig. 7). Etymology The species is named in honour of the author’s husband Richard (Bert) Roberts who collected the first specimens and has become quite expert in the collection of Onychophora. Planipapillus biacinoides, sp. nov. (Figures 2c; 4c; 5d; 7; 10a; 10b; 10c; 11. Table 1) Material examined Holotype: ©, Victoria, beside Livingstone Ck at intersection of Birregun Rd and Upper Livingstone Tk (6.2 km S of intersection of Cassilis Rd and Birregun Rd), 37°05’S 147°36’E, 300 m, 13 Mar 1999, coll. A. Reid and R. Roberts (MV K7285). Paratypes: Victoria, 20 19, data as for holotype (MV K7286). Proc. Linn. Soc. N.s.w., 122. 2000 16 NEW PLANIPAPILLUS (ONYCHOPHORA) Figure 10. Planipapillus biacinoides, sp. novy.: (a) body, paratype male, 0.75 mm HWE, scale bar 0.30 mm; (b) body, holotype male, 1.12 mm HWE, scale bar 0.30 mm; (c) head, holotype male, 1.12 mm HWE, scale bar 0.30 mm. Planipapillus cyclus, sp. nov.: (d) body, holotype male, 0.82 mm HWE, scale bar 0.50 mm; (e) body, paratype female, 0.85 mm HWE, scale bar 0.50 mm; (f) head, holotype male, 0.82 mm HWE, scale bar 0.20 mm. Proc. Linn. Soc. n.s.w., 122. 2000 A. REID 17 Figure 11. Planipapillus biacinoides, sp. nov., modified region of head, holotype male, 1.12 mm HWE, scale bar 0.20 mm. Diagnosis Antennae without tan banding on and between antennal rings 3-4. Ovoid patch of reduced papillae in males with two sclerotised spikes at posterior margin; plicae in 6—7 rows comprise patch; papillae enlarged on each side of patch, each with single sensory bristle. Crural papillae present in both sexes. Posterior accessory glands folded, with short hook distally. Description Measurements HWE males 0.75—0.98—1.12 mm (n=3, Holotype 1.12 mm HWE); HWE female 0.80 mm. Colour pattern Ground colour greyish-blue. Mid-dorsal dark stripe absent; longitudinal light- coloured band (usually tan) along dorsal midline and short, dark, transverse bars or blotches along midline dorsal to oncopods (Fig. 10a) [not visible in dark specimens (Fig. 10b)]; light dorsolateral ovoid patches in line with oncopods [in small specimens only (Fig. 10a)]; laterally with light patches between oncopods (indistinct, some tan-based papillae). Oncopods with light patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present, very pale. Spinous pads greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum, or darker than rest of ventrum. Antennal rings banded, tan or with tan mottle dorsally, or not banded, ground colour; dorsal banding on alternate rings distal to, and including ring five. Antennal rings Distal eight antennal rings with sensory bulbs; sensory pads with 2—3 rows of sensilla (two in small specimens). Eyes EDI males 0.07—0.08—0.08. EDI females 0.06. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in an ovoid-squarish patch posterior to eyes (Fig. 2c). Patch with two short, sclerotised spikes at posterior margin (Figs 10c and 11). Ovoid patch comprising 6-7 rows of plicae; papillae forming patch blunt conical; papillar scales fused, papillae smooth, each with a sensory bristle. Papillae laterally adjacent to patch cylindrical, Proc. Linn. Soc. N.s.w., 122. 2000 18 NEW PLANIPAPILLUS (ONYCHOPHORA) enlarged; 2—3 rows enlarged papillae, innermost row only slightly longer than rest; each enlarged papilla with single bristle. Head (females) Females with no modification of head papillae. Dorsal integument Males and females with 12 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae cylindrical. Male reproductive tract Male genital pad low, conical. Proximal vasa efferentia separate, do not lie parallel before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Fig. 4c). Male glands and gland papillae Crural papillae on ventral side of oncopods 2—3 and 6—14. Papillae shape differs among oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section (oncopods 2—3 and 6-10) or subconical, not divided into distinct basal and distal regions (oncopods 11—14). Crural glands extend into lateral haemocoel from oncopods 11-14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae present; large, semicircular. Anterior accessory glands present; greatly reduced. Posterior accessory glands folded distally, short blunt hook (Fig. 5d). Female crural papillae Present (very small papillae visible on some oncopods, see Remarks). Remarks Planipapillus biacinoides, sp. nov. is very similar to P. biacinaces. The pigmentation of the two species differs. Planipapillus biacinaces specimens have a distinctive broad tan band at the base of each antenna, on and between the third and fourth antennal rings; this trait is not seen in P. biacinoides, sp. nov. This character was not mentioned in the original description of P. biacinaces, but was discovered when comparing P. biacinaces with the species described here. Six to seven plicae comprise the male head patch in P. biacinoides, sp. nov., while 10—11 plicae comprise this patch in P. biacinaces. Planipapillus biacinoides, sp. nov. differs from all other Planipapillus (with the exception of P. biacinaces) in having two sclerotised spikes at the posterior margin of the male head patch. Other differences are shown in Figs 2, 5 and Table 1. This species was found with P. gracilis, sp. nov. While other sympatric peripatopsids are known, in all cases to date where one or more species have been collected at a site, they have belonged to different genera. For example, Planipapillus species often occur with Ooperipatus. This 1s the first time, within Australia that representatives of the same genus have been found together. The heads of males of the two species are distinctly different (compare Figs 10c and 13c), but it is difficult to determine to which species females collected at the site belong. Four females were collected at the site. One of these was very small, but appeared to have crural papillae on some of the oncopods. Female P. biacinaces are now known to have crural papillae on some of the oncopods, so it is highly likely that they also occur in female P. biacinoides, sp. nov. The three larger females that lacked crural papillae have therefore been assigned to P. gracilis, sp. nov. When more, particularly mature, females are collected, this inference can be verified. In addition, the three larger females had a greater number of papillae counted from the mid-dorsal line to the junction of oncopod 10 (20-22 in P. gracilis, sp. nov. and only 12 in P. biacinoides, sp. nov.). Whether this is significant, or simply the result of small versus large females being compared is yet to be determined. No other morphological characters could be found to distinguish the females of the two species but molecular characters may provide additional clues. Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 19 Habitat In and under logs in dry sclerophyll woodland. Specimens were collected beside road in cleared farming area (Fig. 6). Live, hand collected specimens were usually coiled in spiral when first exposed. Distribution This species is known only from the type locality (Fig. 7). Etymology The similarity between this species and P. biacinaces 1s reflected in the species name that is based on biacinaces with the termination ‘-oides’, meaning ‘like’, or ‘resembling the form of’. Planipapillus cyclus, sp. nov. (Figures 2d; 4d; 5e; 7; 10d; 10e; 10f; 12. Table 1) Material examined Holotype: ©} Victoria, 9 km N of Club Terrace, junction of Errinundra Rd and Combienbar Rd, 37°28’S 148°55’E, 130 m, 16 Jun 1999, coll. A. Reid and A. Skates (MV K7287). Paratypes: Victoria, 2 0 29, data as for holotype (MV K7288). Figure 12. Planipapillus cyclus, sp. nov., modified region of head, holotype male, 0.82 mm HWE, scale bar 0.20 mm. Diagnosis Circular patch of reduced papillae posterior to eyes in males with four subequal spikes on the posterior margin; patch surrounded by single row of enlarged semicircular papillae, each with up to four sensory bristles. Crural papillae on oncopods 2—3 and 6—14 in both sexes. Anterior accessory glands present in males, short. Proc. Linn. Soc. N.s.w., 122. 2000 20 NEW PLANIPAPILLUS (ONYCHOPHORA) Figure 13. Planipapillus gracilis, sp. nov.: (a) body, paratype male, (0.92 mm HWE, scale bar 0.30 mm; (b) body, holotype male, 0.95 mm HWE, scale bar 0.30 mm; (c) head, holotype male, 0.95 mm HWE, scale bar 0.20 mm. Planipapillus impacris, sp. nov.: (d) body, paratype female, 1.20 mm HWE, scale bar 0.50 mm; (e) head, holotype male, 1.12 mm HWE, scale bar 0.25 mm. Proc. Linn. Soc. N.S.w., 122. 2000 A. REID 2] Description Measurements HWE males 0.77—0.80—0.82 mm (n=3, Holotype 0.82 mm HWE); HWE; females 0.85—0.88—0.92 mm (n=2). Colour pattern Ground colour tan, brown or greyish-blue. Mid-dorsal dark stripe present (narrow); longitudinal pale ground-coloured band along dorsal midline (Figs 10d and 10e), short, dark, transverse bars, longitudinal bars, or blotches along midline dorsal to oncopods, and laterally with longitudinal light band dorsal to oncopods. Oncopods with light patches at junction with feet. Papillae around anal opening pigmented as for rest of body, or tan. Ventral pigment pale; mottled with darker patches at oncopod bases. Spinous pads tan, or greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum, or tan (in greyish-blue specimens). Antennal rings banded, tan or with tan mottle dorsally, or not banded, ground colour; dorsal banding on alternate rings distal to, and including ring five (for three quarters of antennal length). Antennal rings Uniform width; distal seven antennal rings with sensory bulbs; sensory pads with 2-3 rows of sensilla. Eyes EDI males 0.08—0.08—0.09; EDI females 0.07—0.07—0.08. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in circular patch posterior to eyes (Figs 2d and 10f). Patch with four sclerotised spikes at medio-posterior margin; median pair much smaller than lateral pair, positioned posterior to large spikes (Fig. 12). Ovoid patch comprising 6—7 rows of plicae; papillae forming patch greatly reduced, or absent; papillar scales fused, papillae smooth, median bristles absent. Papillae adjacent to patch semicircular, enlarged; single row of enlarged papillae forming circle, with gap posterior to spikes (Fig. 12); each enlarged papilla with up to four bristles. Head (females) Females with no modification of head papillae. Dorsal integument Males with 12-13-14, females with 13 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae cylindrical. Male reproductive tract Proximal vasa efferentia lying close together, parallel for part of their length before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Fig. 4d). Male glands and gland papillae Crural papillae on ventral side of oncopods 2—3 and 6—14. Papillae similar in shape on all oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section. Crural glands extend into lateral haemocoel from oncopods 6-14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae present; semicircular. Anterior accessory glands present; short. Posterior accessory glands folded distally, long hook tapered only slightly to blunt tip (Fig. Se). Female crural papillae Present on oncopods 2-3 and 6-14. Remarks Planipapillus impacris, sp. nov. males also have four spikes (one large and one small pair) on the posterior margin of the head patch. The head patch is much larger in P. impacris, sp. nov. than in P. cyclus, sp. nov., and is oval, rather than circular in shape. The Proc. Linn. Soc. N.s.w., 122. 2000 i) i) NEW PLANIPAPILLUS (ONYCHOPHORA) papillae surrounding the patch are enlarged to a greater extent in P. impacris, sp. nov. than in P. cyclus, sp. nov., and the spikes at the posterior margin of the patch are larger (compare Figs 12 and 15). Additional characters distinguishing these two species and other Planipapillus species are shown in Figs 2, 5 and Table 1. Habitat The type specimens were found in a log in wet sclerophyll forest. Live, hand collected specimens were usually coiled in spiral when first exposed, the head is tucked in the loop of the body. Distribution This species is known only from the type locality (Fig. 7). Etymology The species name is Latin and means ‘circle’, or ‘ring’. It refers to the shape of the patch of reduced papillae, surrounded by a prominent ring of papillae on the heads of males in this species. Planipapillus gracilis, sp. nov. (Figures 2e; 4e; 5f; 5g; 7; 13a; 13b; 13c; 14. Table 1) Material examined Holotype: ©; Victoria, beside Livingstone Ck at intersection of Birregun Rd and Upper Livingstone Tk (6.2 km S of intersection of Cassilis Rd and Birregun Rd), 37°05’S 147°36’E, 300 m, 13 Mar 1999, coll. A. Reid and R. Roberts (MV K7289). Paratypes: Victoria, 30 39, data as for holotype (MV K7290). Diagnosis Body without median longitudinal light-coloured band. Ovoid patch of reduced papillae on the heads of males without sclerotised spikes; 19—20 rows of plicae comprise patch; 1—2 rows of papillae lateral to patch elongate, finger-like with up to three sensory bristles. Anterior accessory glands absent and gland papillae usually absent. Posterior accessory glands straight, blunt, or tapered to a blunt point distally. Figure 14. Planipapillus gracilis, sp. nov., modified region of head, holotype male, 0.95 mm HWE, scale bar 0.25 mm. Proc. Linn. Soc. n.s.w., 122. 2000 A. REID 23 Description Measurements HWE males 0.90—0.94—1.00 mm (n=4, Holotype 0.95 mm HWE); HWE females 1.07—L.12-1.17 (n=3). Colour pattern Ground colour dark greyish-blue. Mid-dorsal dark stripe absent (Figs 13a and 13b); without median longitudinal light-coloured band; with short, dark, transverse bars or blotches along midline, dorsal to oncopods [trace only visible in dark specimens (Fig. 13b)]; laterally with light patches between oncopods (distinctive). Oncopods with cream patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present; lighter than dorsum. Spinous pads greyish-blue. Integument between genital and anal openings darker than rest of ventrum. Antennal rings banded, tan or with tan mottle dorsally, or not banded, ground colour. Antennal rings Distal 8—9 antennal rings with sensory bulbs; sensory pads with 1—2 rows of sensilla. Eyes EDI males 0.05—0.06—0.06; EDI females 0.05—0.06-—0.07. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in longitudinal ovoid patch posterior to eyes (Figs 2e and 13c). Patch without sclerotised spikes at medio-posterior margin. Ovoid patch comprising 19-20 rows of plicae; papillae forming patch triangular, ridge-like; papillar scales fused, papillae smooth, each with a sensory bristle. Papillae laterally and posteriorly adjacent to patch cylindrical, enlarged; 1-2 rows of enlarged papillae laterally, inner row longest, long and shorter papillae alternate in length from anterior to posterior (Fig. 13c and 14); single row of enlarged papillae posterior to patch; each enlarged papilla with up to three bristles. Head (females) Females with no modification of head papillae. Dorsal integument Males with 12 papillae counted from mid-dorsal line to junction of oncopod 10, females with 20-22-22 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae cylindrical. Male reproductive tract Proximal vasa efferentia separate, do not lie parallel before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Fig. 4e). Male glands and gland papillae Crural papillae on ventral side of oncopods 2—3 and 6—14 (papillae on oncopods 6— 7 reduced). Papillae shape differs among oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section (oncopods 2-3 and 6-9) or subconical, not divided into distinct basal and distal regions (oncopods 10- 14). Crural glands extend into lateral haemocoel from oncopods 11-14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae absent (usually). Anterior accessory glands absent. Posterior accessory glands usually straight, blunt (Fig. 5f), or tapered to blunt point distally (Fig. 5g). Female crural papillae Absent. Remarks The holotype has very reduced anterior accessory papillae on the last oncopod pair, but these papillae are not visible on other material examined. The females (collected in March 1999) contained thick-shelled eggs in the oviducts. Among Planipapillus males that lack spikes at the posterior margin of a large head patch, P. gracilis, sp. nov. is most similar to P. berti. Differences between these two species are Proc. Linn. Soc. N.s.w., 122. 2000 24 NEW PLANIPAPILLUS (ONYCHOPHORA) given in the Remarks section of P. berti. It differs from P. annae, P. bulgensis and P. mundus in having up to three sensory bristles on the enlarged papillae, rather than one on each papilla. The size of the male head patch and elongate papillae surrounding the patch distinguish P. gracilis, sp. nov. from P. bulgensis. The papillae on each side of the patch are much more elongate in P. gracilis, sp. nov. than they are in P. annae and P. bulgensis. Additional characters that distinguish P. gracilis, sp. nov. from other Planipapillus species are given in Figs 2, 5 and Table 1. Planipapillus gracilis, sp. nov. was found with Planipapillus biacinoides. Females of these two species may be difficult to distinguish using morphological characters (see the Remarks section of P. biacinoides above) but the differences between the heads of males are very obvious (compare Fig. 10c with Fig. 13c). Habitat Planipapillus gracilis, sp. nov. was found in and under dry pieces of timber beside Livingstone Ck. Though largely cleared, this habitat was once open dry sclerophyll forest. Live, hand collected specimens were usually coiled in spiral when first exposed. Distribution This species is known only from the type locality (Fig. 7). Etymology The Latin specific name ‘gracilis’, means slender, or thin. Planipapillus impacris, sp. nov. (Figures 2f; 4f; 5h; 7; 13d; 13e; 15; 16. Table 1) Material examined Holotype: 0; New South Wales: South East Forests NP, Coolangubra Section, 5 km N of intersection of Coolangubra Forest Way and Northern Access Rd, 37°01’S 149°23’E, 800 m, 2 Mar 1999, coll. A. Reid (MV K7291). Paratypes: 1 0; 40 , 1 juvenile, data as for holotype (MV K7292). Additional material. 10°, South East Forests NP, Coolangubra Section, Waratah Forest Rd., 2 Feb 1989, coll. R. Cameron (MV). Figure 15. Planipapillus impacris, sp. noyv., modified region of head, holotype male, 1.12 mm HWE, scale bar 0.25 mm. Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 25 Figure 16. Planipapillus impacris, sp. nov., male reproductive tract and associated glands, holotype, 1.12 mm HWE, scale bar 0.50 mm. aag, anterior accessory gland; cg, crural gland; pa, posterior accessory gland; sv, seminal vesicle; t, testis; vd, vas deferens; ve, vas efferens. Diagnosis Ovoid patch of reduced papillae in males with four subequal spikes on posterior margin; patch surrounded by single complete row and second partial row (with one dorsolateral and one ventrolateral pair of enlarged papillae) of semicircular papillae; each enlarged papilla with up to seven sensory bristles. Crural papillae on oncopods 6—14 in both sexes. Anterior accessory glands present, greatly reduced. Proc. Linn. Soc. N.S.W., 122. 2000 26 NEW PLANIPAPILLUS (ONYCHOPHORA) Description Measurements HWE males 0.82—0.98—1.12 mm (n=3, Holotype 1.12 mm); HWE females 1.05— 1.12—1.20 mm (n=4). Colour pattern Ground colour greyish-blue. Mid-dorsal dark stripe present (Fig. 13d); longitudinal light-coloured band along dorsal midline (juvenile only) and short, dark, transverse bars or blotches along midline, dorsal to oncopods; tan dorsolateral ovoid patches in line with oncopods (juvenile only), or irregular mottling; tan papillae sometimes concentrated mediodorsally (Fig. 13d); laterally with cream or tan patches between oncopods. Oncopods without light patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present, very pale. Spinous pads greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum. Antennal rings banded, tan or with tan mottle dorsally, or not banded, ground colour; dorsal banding on proximal half of each antennal ring (distal half ground colour), with every fourth ring predominantly tan. Antennal rings Distal 7—9 antennal rings with sensory bulbs; sensory pads with 2-3 rows of sensilla. Eyes EDI males 0.06—0.06—0.07; EDI females 0.06—0.06—0.07. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in transverse ovoid patch posterior to eyes (Figs 2f and 13e). Patch with four sclerotised spikes at medio-posterior margin; median pair smaller than lateral pair; median pair slightly posterior to lateral pair, curved (Fig. 15). Ovoid patch comprising 6— 7 rows of plicae; papillae forming patch greatly reduced, or absent; papillar scales fused, papillae smooth, median bristles absent. Papillae adjacent to patch semicircular, enlarged; single row of enlarged papillae, except for one enlarged dorsolateral and one enlarged ventrolateral pair in outer row. Papillae posterior to median two spikes smaller than rest; each enlarged papilla with up to 7 bristles. Head (females) Females with no modification of head papillae. Dorsal integument Males with 11-15-17, females with 16-17-18 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae semicircular. Male reproductive tract Male genital pad cylindrical, protuberant. Proximal vasa efferentia lying close together, parallel for part of their length before fusing to form vas deferens, or separate, do not lie parallel for part of their length (Figs 4f and 16); vas deferens continues directly (without looping) from paired vasa efferentia to gonopore. Male glands and gland papillae Crural papillae on ventral side of oncopods 6—14. Papillae similar in shape on all oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section (papillae on oncopods 10-14 broader than rest). Crural glands extend into lateral haemocoel from oncopods 9-14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae present; large, subconical. Anterior accessory glands present; greatly reduced (Fig. 16). Posterior accessory glands with long hook tapered only slightly to blunt tip (Figs 5h and 16). Female crural papillae Present on oncopods 6—14. Not visible on all specimens. Proc. Linn. Soc. N.S.w., 122. 2000 A. REID 27 Remarks The head spikes are well-developed in a small male, 0.82 mm HWE. Planipapillus impacris, sp. nov. differs from P. cyclus (also with four subequal spikes in the posterior margin of the head patch in males) in the absence of crural papillae on oncopods 2-3 in both sexes. Planipapillus cyclus has crural papillae on oncopods 2-3. The head spikes also differ between the two species. The median pair are much larger, and distinctly curved in P. impacris, sp. nov. The patch of modified papillae is ovoid in P. impacris, sp. nov., and circular in P. cyclus (compare Figs 10f and 13e). Differences between P. impacris, sp. nov. and other Planipapillus are given in Figs 2, 5 and Table 1. This species was described as ‘taxon I’ by Tait and Briscoe (1990). Planipapillus impacris, sp. nov. was found with specimens of Ooperipatus sp. Habitat In decomposing logs. Live, hand collected specimens were usually coiled in spiral when first exposed. Distribution Known only from the type locality (Fig. 7). Etymology The specific name is derived from the Latin ‘impar’, meaning unequal, or odd, and ‘acris’, meaning pointed. The name refers to the unequal length of the two pairs of head spikes in this species. Planipapillus tectus, sp. nov. (Figures 2g; 4g; 51; 5j; 5k; 7; 17a; 17b. Table 1) Material examined Holotype: ©; Victoria, 6.7 km S of intersection of Gelantipy Rd and Tulloch Ard Rd (10.7 km S of Gelantipy, 300 m N of Forest Ck Tk), 37°17’S 148°15’E, 710 m, 14 Mar 1999, coll. A. Reid and R. Roberts (MV K7293). Paratypes: Victoria, 30/29, data as for holotype, (MV K7294). Additional material: Victoria, 29, Gelantipy, Honeysuckle Tk, Apr 30—May 9 1947, coll. C.W.B and Miss M.B. (MV). Diagnosis Males with ovoid patch of slightly reduced papillae on head; anteriorly papillae within patch semicircular with scales; posteriorly papillae ridge-like, smooth, with fused scales; papillae surrounding patch not markedly enlarged. Crural papillae on oncopods 2-3 and 6—14 in both sexes. Anterior accessory glands greatly reduced. Posterior accessory glands straight, blunt (usually), or folded with long hook. Description Measurements HWE males 0.80—0.82—0.85 mm (n=4, Holotype 0.85 mm HWE); HWE females 0.80—0.85—0.90 mm (n=2). Colour pattern Ground colour greyish-blue. Mid-dorsal dark stripe absent (Fig. 17a); longitudinal light ground-coloured band along dorsal midline (Fig. 17a); short, dark, transverse bars or blotches along midline dorsal to oncopods and light ground colour dorsolateral transverse patches in line with oncopods, patches flat anteriorly, convex posteriorly, often with prominent whitish papilla on lateral margin of each patch (Fig. 17a), or irregular Proc. Linn. Soc. N.s.w., 122. 2000 28 NEW PLANIPAPILLUS (ONYCHOPHORA) mottling (one small specimen 0.8 mm HWE); laterally with longitudinal light band dorsal to oncopods, or with light patches between oncopods. Oncopods with cream patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present; ground colour, lighter than dorsum. Spinous pads greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum. Antennal rings banded, tan or with tan mottle dorsally; dorsal banding on alternate rings distal to, and including ring five (every 5th ring with more tan mottle). Antennal rings Distal 8—9 antennal rings with sensory bulbs; sensory pads with two rows of sensilla. Eyes EDI males 0.06—0.07—0.08; EDI females 0.07. Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in longitudinal ovoid patch posterior to eyes (Figs 2g and 17b). Patch without sclerotised spikes at medio-posterior margin. Ovoid patch comprising 15-17 rows of plicae; papillae forming patch only very slightly reduced, semicircular anteriorly, posteriorly papillae are triangular, ridge-like (Fig. 17b); anterior semicircular papillae with scales as for remaining head papillae, not fused; posterior ridge-like papillar scales fused, papillae smooth; each with a sensory bristle. Papillae laterally adjacent to patch semicircular, very slightly enlarged; each enlarged papilla with single bristle. Head (females) Females with no modification of head papillae. Dorsal integument Males and females with 12 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae cylindrical. Male reproductive tract Male genital pad semicircular. Proximal vasa efferentia separate, do not lie parallel before fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Fig. 4g). Male glands and gland papillae Crural papillae on ventral side of oncopods 2—3 and 6-14. Papillae shape differs among oncopods: oncopods 2-3 cylindrical, oncopods 6—10 and 11-14 papillae semicircular; papillae tapered slightly distally, not divided into distinct basal and distal regions. Crural glands extend into lateral haemocoel from oncopods 11—14; straight, short, not folded; remaining glands confined within oncopods. Anterior accessory gland papillae present; broad, rounded. Anterior accessory glands present; greatly reduced. Posterior accessory glands variable: usually straight, bulbous, blunt distally (Fig. 51); folded distally with short blunt hook (Fig. 5j); or sometimes constricted forming distal knob (Fig. 5k). Female crural papillae Females with crural papillae on oncopods 2-3 and 6-14. Remarks Unlike other Planipapillus, the papillae on the head patches of male P. tectus, sp. noy. are only slightly reduced. Some papillae have ribbed scales and sensory bristles, while others are reduced and ridge-like with fused scales within the head patch. In other Planipapillus the papillae are fairly uniformly modified within the head patch. The papillae lateral to the patch in P. tectus, sp. nov. are only slightly elongate, thus differing from most other Planipapillus, with the exception of P. vittatus, sp. nov. Additional characters that distinguish P. tectus, sp. nov. from other nominal Planipapillus are shown in Figs 2, 5 and Table 1. The Additional material, two female specimens, are only tentatively assigned to this species. One of these specimens is a large female (1.25 mm HWE) and differs from Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 29 Figure 17. Planipapillus tectus, sp. nov.: (a) body, paratype female, 0.90 mm HWE scale bar 0.20 mm; (b) head, holotype male, 0.85 mm HWE, scale bar 0.20 mm. Planipapillus vittatus, sp. nov. (the position of the patch of modified papillae is indicated by an arrow): (c) body, paratype female, 1.0 mm HWE, scale bar 0.50 mm; (d) head, paratype male, 0.90 mm HWE, scale bar 0.20 mm. the type material in being of tan colouration, rather than greyish-blue, although the pattern is the same as the other material examined. This specimen, collected in May 1947, contains thick-shelled eggs in the oviducts. Habitat Under logs in dry sclerophyll forest. Live, hand collected specimens were usually coiled in spiral when first exposed. Proc. Linn. Soc. N.s.w., 122. 2000 30 NEW PLANIPAPILLUS (ONYCHOPHORA) Figure 18. Planipapillus vittatus, sp. nov., modified region of head, holotype male, 0.85 mm HWE, scale bar 0.20 mm. Distribution This species is known only from the type locality (Fig. 7). Etymology The species name is Latin, meaning ‘cover’. It refers to the behaviour of this species, found under the cover of logs. Planipapillus vittatus, sp. nov. (Figures 4h; 41; 51; 7; 2h; 17c—d; 18. Table 1) Material examined Holotype: ©; Victoria, Dinner Plain, 36°59’S 147°17’E, 1628 m, 12 Mar 1999, coll. A. Reid and R. Roberts (MV K7295). Paratypes: Victoria, 20% Ts data as for holotype (MV K7296); 29; Dinner Plain, 36°59’S 147°17’E, 11 Jan 1990, coll. N.N. Tait (MV K7297). Diagnosis Patterned with distinctive median longitudinal tan stripe on dark grey body. Males with small patch of reduced ridge-like papillae on head; 5—6 rows plicae comprise patch; median pair largest; papillae surrounding patch not enlarged. Males with crural papillae on oncopods 2-3 and 6-14; females with crural papillae on oncopods 6-14. Description Measurements HWE males 0.77—-0.84—0.90 mm (n=3, Holotype 0.85 mm HWE); HWE females 0.57— 0.86—1.00 mm (n=9). Colour pattern Ground colour dark greyish-blue (sometimes with tan mottle). Mid-dorsal dark stripe absent; distinctive longitudinal tan band along dorsal midline, extending from midpoint of eyes to last oncopod pair (Figs 17c and 17d), band sometimes interrupted by greyish-blue or brown patches dorsal to oncopods; evenly scattered tan or tan-based papillae (some speci- mens only); cream dorsolateral transverse patches in line with oncopods [juvenile (0.57 mm HWE) only]; laterally with longitudinal cream band dorsal to oncopods, or with light patches between oncopods. Oncopods without light patches at junction with feet. Papillae around anal opening pigmented as for rest of body. Ventral pigment present, very pale. Spinous pads greyish-blue. Integument between genital and anal openings pigmented as for rest of ventrum. Proc. Linn. Soc. N.s.w., 122. 2000 A. REID 31 Antennal rings not banded, ground colour. Antennal rings Distal 8—9 antennal rings with sensory bulbs; sensory pads with two rows of sensilla. Eyes EDI males 0.05—0.05—0.06 (n=3); EDI females 0.04—0.05—0.07 (n=9). Head (males) Males with modified papillae on head (i.e. differ from remaining dorsal papillae). Papillae reduced in an ovoid-squarish patch posterior to eyes (Fig. 2h). Patch without sclerotisea spikes at medio-posterior margin (Figs 17d and 18). Ovoid patch comprising 5-6 rows of plicae; papillae forming patch triangular, ridge-like; papillar scales fused, papillae smooth; 2—4 modified papillae on each plica, median pair largest (Figs 17d and 18), median bristles absent. Papillae adjacent to patch not enlarged; each papilla with up to two bristles. Head (females) Females with no modification of head papillae. Dorsal integument Plicae uniform width. Males with 10—10—11, females with 6-12-17 papillae counted from mid-dorsal line to junction of oncopod 10. Primary papillae semicircular. Male reproductive tract Proximal vasa efferentia lying close together, parallel for part of their length fusing to form vas deferens; vas deferens continues directly (without looping) from paired vasa efferentia to gonopore (Figs 4h and 41). Male glands and gland papillae Crural papillae on ventral side of oncopods 2—3 and 6—14. Papillae similar in shape on all oncopods: semicircular or cylindrical proximally, tapered abruptly to narrower, semicircular or cylindrical distal section (oncopods 2-3) or semicircular or cylindrical, tapered slightly distally, not divided into distinct basal and distal regions (oncopods 6— 14). Crural glands extend into lateral haemocoel from oncopods 11—14; sometimes folded back along length; remaining glands confined within oncopods. Anterior accessory gland papillae present; semicircular. Anterior accessory glands present; greatly reduced. Posterior accessory glands folded distally, long hook tapered only slightly to blunt tip (Fig. 51). Female crural papillae Present on oncopods 6-14. Remarks The bold longitudinal tan stripe on the body distinguishes both sexes of P. vittatus, sp. nov. from all other Planipapillus. The small patch of ridge-like papillae on the heads of males, the median pair of which is enlarged, is also very distinctive in this species. Other distinguishing characters are shown in Figs 2, 5 and Table 1. One female specimen (collected on 11 Jan 1990) has a patch of reduced papillae on the head. This patch of reduced papillae is much larger than the modified patch seen in the males of this species. The region may be the result of a wound to the head, though it is strange that this patch of reduced papillae is in the same position to that seen in males. No other females showed any modification of the head papillae. Females collected on 12 Mar 1999 contained eggs at various stages of development along the length of the oviducts, though none were well-developed. The females collected on 11 Jan 1990 contained large, thick shelled eggs in the oviducts. Habitat Under dry logs at the edge of grassy paddock. The predominant vegetation in the region is snow gum woodland. Live, hand collected specimens were usually lying flat and straight when first exposed. Proc. Linn. Soc. N.s.w., 122. 2000 32 NEW PLANIPAPILLUS (ONYCHOPHORA) Distribution Known only from the type locality (Fig. 7). Etymology The species name, ‘vittatus’ is derived from the Latin, “vitta’, meaning band, or stripe, and refers to the distinctive dorsal pigmentation in this species. ACKNOWLEDGEMENTS I wish to thank Bert Roberts and Anne Skates for their company and assistance with fieldwork. Thanks to Sue Boyd, Chris Rowley and Ken Walker for the use of microscopes and photographic equipment in their departments at Museum Victoria. Many thanks also to Noel Tait for his thorough review of this manuscript and constructive suggestions which resulted in its considerable improvement. In addition, he first discovered three of the sites from which I collected specimens described in this paper. The Australian Biological Resources Study funded this work. I am very grateful for this support. REFERENCES Briscoe, D.A. and Tait, N.N. (1993). Peripatus or velvet worms. In “Tasmanian Wilderness — World Heritage Values’ (Eds S.J. Smith and M.R. Banks) pp.136—138. (Royal Society of Tasmania: Hobart). Dallwitz, M.J. (1980). A general system for coding taxonomic descriptions. Taxon 29, 41-46. Dallwitz, M.J., Paine, T.A. and Zurcher, E.J. (1993). “Users Guide to the DELTA System: a General System for Processing Taxonomic Descriptions’. 4th Edn. (CSIRO Division of Entomology: Canberra.). Dendy, A. (1900). Preliminary note on a proposed new genus of Onychophora. Zoologischer Anzeiger 23, 415-416. Gleeson. D.M. (1996). Onychophora of New Zealand; past present and future. New Zealand Entomologist 19, 51-55. Meredith, P. (1995). Headfirst. Passion goes to the velvet worm’s head. BBC Wildlife 13(6), 26. Partridge, T.R., Dallwitz, M.J. and Watson, L. (1993). ‘A Primer for the DELTA System’. 3rd Edn. (CSIRO Division of Entomology: Canberra.). Reid, A.L. (1996). Review of the Peripatopsidae (Onychophora) in Australia, with comments on peripatopsid relationships. Invertebrate Taxonomy 10(4), 663-936. Sunnucks, P. and Wilson, A.C.C. (1999). Microsatellite markers for the onychophoran Euperipatoides rowelli. Molecular Ecology 8, 899-200. Tait, N.N. and Briscoe, D.A. (1990). Sexual head structures in the Onychophora: unique modifications for sperm transfer 24, 1517-1527. Tait, N.N. and Briscoe, D.A. (1995). Genetic differentiation within New Zealand Onychophora and their relationships to the Australian fauna. Zoological Journal of the Linnean Society 114, 103-113. Trewick, S.A. (1998). Sympatric cryptic species in New Zealand Onychophora. Biological Journal of the Linnean Society 63, 307-329. Proc. Linn. Soc. N.s.w., 122. 2000 Pentremites australis sp. nov., a New Lower Carboniferous (Tournaisian) Blastoid from New South Wales I.D. LINDLEY Department of Geology, Australian National University, Canberra, ACT. 0200. Lindley, I.D. (2000). Pentremites australis sp. nov., anew Lower Carboniferous (Tournaisian) blastoid from New South Wales. Proceedings of the Linnean Society of New South Wales 122, 33-42. A new species of the blastoid Pentremites Say is described from the Lower Carboniferous Brushy Hill Limestone Member (middle Tournaisian) at Glenbawn Dam, New South Wales. Pentremites australis sp. nov. has a small pyriform (obconical) theca strongly pentagonal in plan view, with high pelvis and truncate vault. The anispiracle is excavated in an apparently undivided anal deltoid plate. Its short deltoids are barely visible and lancets are exposed full width forming petaloid ambulacra on the summit. Side plates abut the lancet and one hydrospire pore is present per side plate. Stratigraphically, P. australis is the earliest member of the genus, otherwise known from Mississippian (late Tournaisian) to Lower Pennsylvanian sequences of North and South America. The theca of the new species exhibits a combination of primitive and derived features. Manuscript received 3 July 2000, accepted for publication 18 October 2000. Keywords: Pentremites, Blastoidea, Lower Carboniferous, Tournaisian, New South Wales. INTRODUCTION Blastoids are a rare component of invertebrate faunas in the Carboniferous sequences of Australia and few have been described. Nymphaeoblastus bancroftensis McKellar, 1964, a fissiculate blastoid, was described by McKellar (1964) from the Lower Carboniferous Tellebang Formation, east of Monto, Queensland, in association with a prolific late Viséan Rhipidomella fortimuscula Zone brachiopod fauna. McKellar (1966) reviewed Etheridge’s (1892) specimens from which he described the spiraculate Malchiblastus australis (Etheridge, 1892) from the Late Carboniferous Neerkol Formation. Campbell (1961) described a solitary radial plate from the Late Carboniferous Booral Formation, north of Newcastle, New South Wales, which he tentatively referred to Pentremites. Both the Neerkol and Booral Formations contain Namurian brachiopod faunas of the Levipustula levis Zone. Pentremites Say, 1820 is noted for its relatively long stratigraphic range from the Mississippian (Osagean-Meramecian) to the Lower Pennsylvanian (Morrowan) and geographic distribution extending across North America into South America. Pentremites and the Pentremitidae have been the subject of numerous taxonomic revisions (Galloway and Kaska 1957; Macurda 1975; Macurda and Breimer 1977; Horowitz et al. 1981; Waters et al. 1985; Horowitz et al. 1986; Waters and Horowitz 1993) and eighteen species of Pentremites are presently known. Pentremites kirki Hambach, 1903 and Pentremites elongatus Shumard, 1855, in the Osagean (late Tournaisian - Tn 3: Jones, 1996) Burlington Limestone and its equivalents in North America, represent the earliest species of the genus (Waters et al. 1985). Proc. Linn. Soc. N.s.w., 122. 2000 LOWER CARBONIFEROUS BLASTOID This paper describing Pentremites australis sp. nov. from the Lower Carboniferous Brushy Hill Limestone Member at Glenbawn Dam, in the Hunter Valley of New South Wales, provides the first record of the genus outside the Americas and the oldest record of the genus. The limestone occurs near the base of the Dangarfield Formation, an 850 m thick sequence consisting of mudstone, siltstone, sandstone and calcarenite. Crinoids are locally prolific in calcarenites in the upper Dangarfield Formation, and include camerates and platycrinitids. Some of the camerate crinoids have been described by Lindley (1979, 1988). Terminology used herein follows that of Beaver (1967) and Waters et al. (1985). STRATIGRAPHY The specimen of Pentremites australis was collected by J. Roberts from a quarry in the Brushy Hill Limestone Member near the southern abutment of Glenbawn Dam. Subsequent engineering works on the dam have resulted in changes to the quarry. A [_] attuvium Se = Dangarfield Formation YS = oe mudstone & minor lithic - cE a =) GLENBAWN DAM 46 — sandstone, limestone = — Brushy Hill Limestone Member oolitic limestone Macqueen Formation = \ lithic sandstone, siltstone yy & mudstone = \ " Kingsfield Formation 45 crystal tuff \ SS \ A= TO MUSWELLBROOK \ = 0 1000m \ = aero eee a \ 44 -— G Quarry \ | = : F z= SS @ Collection locality ( 265mm), cobbles (64-256mm), pebbles (4- 64mm), gravel (2-4mm), and sand (<2mm) (Cummins, 1962). The proportion of each substrate size class in each lake was estimated by walking the perimeter with a tape mea- sure. Sites dominated by boulders and/or cobbles were chosen. The number of sites cho- sen for each lake varied to represent the size of the lake (Albina — Al to A4, Blue — B1 to B5, Club — Cl to C3, Cootapatamba — D1 to D3) (Fig. 1). All sites were sampled in December, but in March only one site per lake was worked (A2, B1, C1, D1). At each site the relative proportions of different particle sizes were estimated by dropping randomly a | m? quadrat three times and averaging the perceived proportions of the Wentworth size classes mentioned above by three investigators. The fauna was then sampled by two methods, sweeping and cobble picking. Sweeping involved investigators disturbing a strip of habitat 1m x 10m with their feet and sweeping the disturbed water with a pond net of aperture 700 cm? and mesh size of 0.5 mm for one minute. For the cobble picking method, 10 cobbles per site were selected. These were approximately 10- 20 cm diameter and were removed one by one with the net underneath to catch escaping organisms. The cobbles and the net contents were then placed on a tray and the organisms removed. The bottom area of each rock was estimated by measuring two perpendicular diameters and the average diameter used to calculate area. In both methods, the organisms caught were preserved in 70% ethanol for later identification, enumeration and weighing. Oligochaetes and platyhelminthes were counted in the field due to individuals in these groups breaking up on preservation, although they were still included in collections for biomass measurements. Biomasses were estimated by blotted wet weights using a Sarto- rius top-loading electronic balance (+0.001 g). The shells of molluscs and cases of caddis fly larvae were removed prior to weighing. Proc. Linn. Soc. N.s.w., 122. 2000 M.A. HANCOCK, B.V. TIMMS, J.K. MORTON AND B.A. RENSHAW 71 Lake Cootapatamba D1 Lake Albina C2 Club Lake Blue Lake Figure 1. Sampling sites in the Alpine lakes of the Kosciuszko region (Modified from Dulhunty, 1945). Scale bar applies to all lakes. Major creeks indicated — outflows are distinguished with an arrow. Statistical Methods Similarities in macroinvertebrate assemblage composition among lakes and seasons were examined using a non-metric multidimensional scaling (NMDS) ordination. Bray- Curtis dissimilarities were calculated for each pair of samples on abundances of all spe- cies. To ordinate the data, the subroutine MDS in PRIMER was used, employing 99 ran- dom starts to minimise the risk of erroneously accepting solutions trapped in local minima (Clarke and Warwick 1994). The ANOSIM (Clarke and Warwick 1994) procedure was used to test for differences between lakes. Environmental variables were correlated with ordination scores using Primary Axis Correlation (PCC) in PATN. The relationship between sweep sample biomasses and substrate type was examined using a Pearson correlation. Substrate type was expressed as a substrate index, calculated by ranking each rock size class (boulders=1, cobbles=2, pebbles=3, gravel=4, sand=5) and then calculating a weighted average based on the percentages of each size class in the quadrats. Proc. Linn. Soc. N.s.w., 122. 2000 72 INVERTEBRATE COMMUNITIES OF KOSCIUSZKO LAKES RESULTS Although each of the lakes has a rocky littoral zone, the proportion of rock size classes varies among sites (Table 1). Blue Lake is dominated by bedrock and boulders in a littoral zone that is narrow and shelves deeply and in Club Lake cobbles and pebbles dominate a broad littoral. One side of Lake Cootapatamba is dominated by boulders in deeply shelving water and the other side by cobbles and pebbles in a broad littoral. Habitat complexity is greatest in Lake Albina, aided further by its much greater relative shoreline length, i.e. shoreline development (see Timms 1992). Table 1 Proportion of dominant littoral substrate types around each lake. Substrate index range for each lake. Substrate index explained in text. Particle sizes classified according to the Wentworth Scale (Cummins 1962). [Stubstrately pe unaaaat ania A lb iniallenINnnai nS | uc mannan Unni nnE Club aE Cootapatanibaamm % % % % Bedrock 4 24 2 4 Boulders 27 61 15 42 Cobbles 38 14 44 22 Pebbles 19 0 26 15 Gravel 12 0 13 17 Substrate Index 1.6-2.8 0.9-2.6 2.6-2.9 1.9-3.1 Thirty six taxa of macroinvertebrates were found in the four lakes (Table 2). Species richness was considerably greater in Lake Albina than in the other lakes and was higher in sweeps than in cobble picks. There was a decrease in species richness between early sum- mer and autumn, probably associated with breeding cycles and insect emergence. Many insects were in late instars in December and apparently absent in March, while crusta- ceans bred early as judged by the many juveniles and females with larvae but not eggs in December. Flatworms and tubificids were widespread but not common and probably not speciose. Molluscs and crustaceans were also of limited diversity but were common, often domi- nant and widespread (Table 2). A bivalve, a snail (the endemic Glacidorbis hedleyi), a phreatoicid isopod and an amphipod (probably endemic) occurred in almost all lakes, while Blue Lake had a further snail of widespread distribution in Australia, Austropeplea tomentosa. By contrast insects were diverse, but distribution was patchy and typically they were not abundant (Table 2). Mayflies were common in only Lakes Albina and Cootapatamba, stoneflies (particularly Eusthenia venosa) in Blue Lake, chironomids in Albina and Blue Lakes, and beetles in Lake Albina. Only trichopterans were relatively common and shared between the lakes, with an unidentified limnephid caddis widespread and often abundant. Biomass of sweep samples suggested Albina and Club Lakes had the highest stand- ing crops and Blue Lake by far the lowest (Fig. 2). However, the cobble picking method gave a different order: Club and Blue Lakes the highest, with Albina and Cootapatamba Lakes with about half as much (Fig. 2). Contributions by major taxa to these totals also varied with method and within and between lakes (Fig. 2). Crustaceans (mainly Metaphreatoicus australis) usually domi- nated in sweeps, with molluscs (mainly Psidium kosciusko) important in Club Lake, and insects (mainly mayflies) of some importance in Lake Albina. The cobble picking method gave a different result — Molluscs hardly featured at all (except in Blue Lake), crustaceans Proc. Linn. Soc. n.s.w., 122. 2000 M.A. HANCOCK, B.V. TIMMS, J.K. MORTON AND B.A. RENSHAW Table 2 Littoral species in the Kosciuszko lakes. S = mean number of individuals per m’ of sweep in each lake (bold) ; C = mean number of individuals per m? of cobble in each lake. Albina Blue Club Cootapatamba Species S C S Cc S Cc S C Platyhelminthes Unidentified planarians 1.5 21.1 1.1 0.1 14.3 Sop Annelida: Oligochaeta Unidentified tubificids 0.2 3.5 0.2 1.4 0.3 2.9 0.3 1.5 Mollusca: Bilvalva Pisidium kosciusko 10.7 1.3 1.4 194.6 141.1 13.0 0.2 Mollusca: Gastropoda Glacidorbis hedleyi 5.4 10.6 1.4 8.4 3219 Austropeplea tomentosa 2.6 6.4 Crustacea: Isopoda Metaphreatoicus australis 32.1 61.6 0.4 353 82.2 43.1 16.8 19.6 Crustacea: Amphipoda Neoniphragus n.sp. 0.3 0.9 0.4 2.9 11.0 128.0 Insecta: Ephemeroptera Tasmanophlebia lacascoerulei 0.9 3.6 0.1 Ameletoides lacusalbinae 7 20:6 0.2 1.5 Nousia sp. and Tillyardophlebia alpina 0.2 7.8 0.1 0.5 SES) Insecta: Plecoptera Eusthenia venosa 0.6 Datel Notonemouridae nymph 0.1 0.1 0.2 5.8 <0.05 Insecta: Hemiptera Sigara sp. <0.05 Insecta: Mecoptera Insecta: Coleoptera Nannochorista sp. 0.1 Insecta: Trichoptera Ecnomus sp. 0.2 eS Plectrocnemia sp. 0.3 4.3 0.1 5.4 Austrorheithrus sp. 0.1 <0.05 3.1 <0.05 <0.05 Limnephidae larvae 0.9 <0.05 21.2 0.6 62.3 38.3 Leptoceridae larvae 0.1 0.1 Insecta: Diptera Procladius villosimanus 0.5 4.0 Tanytarsus sp. 0.3 Paramerina levidensis? 0.0 2.6 Botryocladius grapeth 0.1 Polypedilum sp. 0.2 1.7 0.1 7.8 0.1 0.1 Unidentified ceratopogonid <0.05 Tipulidae sp 1 <0.05 Tipulidae sp 2 <0.05 Antiporus femoralis adults <0.05 Sternopriscus wehnekei adults 0.2 1.3 Sternopriscus larvae 2.6 Ds) Elmidae sp 1 adults 0.2 So) Elmidae sp 2 adults 0.2 1.8 1.0 Elmidae larvae 0.2 Curculionidae adults 0.1 0.1 Sclerocyphon basicollis larvae 0.0 2.0 Scirtidae larvae <0.05 Species Richness 25.0 19.0 12.0 12.0 15.0 7.0 14.0 9.0 Total Species Richness 26.0 15.0 15.0 17.0 Proc. Linn. Soc. N.S.w., 122. 2000 74 INVERTEBRATE COMMUNITIES OF KOSCIUSZKO LAKES were sometimes important (in Lakes Albina and Cootapatamba) but insects usually domi- nated (mayflies, caddis and beetles in Lake Albina, large stoneflies and caddis in Blue Lake, caddis in Club Lake and Lake Cootapatamba). 12 10 Biomass (S - g/10m2, C - g/m”) n 4 2 0 1 Sie SS Ss C Ss ¢€ Albina Blue Club Cootapatamba Figure 2. Mean total biomass for Lakes. S indicates sweep samples and C cobble samples. Standard error bars shown. Proportion of biomass consisting of insects (black), crustaceans (open) and molluscs (hatched). All other invertebrates <0.09 and not shown. Biomass figures also changed with season. Sweep biomasses declined by a factor of 1.3 to 20 between December and March, but figures for cobble picks were equivocal, declining by a factor of 2.7 and 5 in Club and Blue Lakes respectively, but increasing by a factor of 1.2 and 5 in Cootapatamba and Albina respectively. Decreases in the figures from sweeps was due mainly to insects, but the increase in the cobble pick figures was due mainly to crustaceans. 4 0 1 2 -2 1 0 1 2 Figure 3. Ordination of littoral assemblages in lakes Albina (circles), Blue (triangles), Club (squares) and Cootapatamba (diamonds). December samples (open symbols) and March samples (closed symbols). (a) Sweep samples, stress = 0.11. (b) Cobble pick samples, stress = 0.14. Sites within each lake most dominated by boulders (high substrate index) are underlined. Proc. Linn. Soc. n.s.w., 122. 2000 M.A. HANCOCK, B.V. TIMMS, J.K. MORTON AND B.A. RENSHAW 75 Ordination of both sweep and cobble pick samples demonstrate that sites group by lake (Fig. 3). Pairwise ANOSIM tests show that the four lakes are significantly different (all pairwise tests significant PS0.05, Global R=0.440, 0.006 M8 Dimtscmal Morten puma, WidIte Aerie Tae prletyenient 0, ch ieee Pyoeetanane ead. teh ( havin the Pallavi (neon vols Ay ied hay fee Perv AY an me f Perle ity, “xia” Poder £ oath riionounttae, disc TNA Og de PLT Vp eel hom TD name Cetrigt Wie Tie lee. Wittiince: Rie se | yi vei Cee Wutie = Vi 2 Toocks. Wie qpeeciad(® Aiveak Agidecw A euties And Sela Mbachesls: for aanie : { A i fixe ride Ly Pw Ue iasinive HyV : Mr sees n.4e ae H ; , Autotrophic Picoplankton in a Regulated Coastal River in New South Wales TsuyosH! KoBaAyASHt, SIMON WILLIAMS? AND AMANDA KoTLASH® 'AWT Environment Science and Technology, PO Box 73, West Ryde NSW 2114 and Centre for Marine and Coastal Studies, University of New South Wales, Sydney NSW 2052 (present address: NSW Environment Protection Authority, PO Box A290, Sydney South NSW 1232); 2 AWT Environment Science and Technology, PO Box 73, West Ryde NSW 2114 (present address: Sydney South Coast Region, Department of Land and Water Conservation, PO Box 867, Wollongong NSW 2520); and 3AWT Environment Science and Technology, PO Box 73, West Ryde NSW 2114 (present address: Blue Mountain City Council, PO Box 189, Katoomba NSW 2380) Kobayashi, T., Williams, S. and Kotlash, A. (2000). Autotrophic picoplankton in a regulated coastal river in New South Wales. Proceedings of the Linnean Society of New South Wales 122, 79-88. Cell density, cell type and vertical distribution of autotrophic picoplankton (APP, cell size 0.2- 2 um) were examined for a year from September 1992 at three freshwater sites in the Hawkesbury-Nepean River. During the study, mean cell density of APP at 1 m deep varied seasonally two orders of magnitude from 2.2 x 10° to 3.2 x 10° cells mL”. At upstream sites of Penrith and North Richmond, higher cell density was observed from summer to autumn. There were three cell types of APP (i.e. coccoid, ellipsoid and rod-shaped). Proportionally, coccoid cells increased downstream from 25 to 52 % of total cells, whereas ellipsoid and rod-shaped cells decreased downstream from 64 to 47 % and from 11 to 1 % of total cells, respectively. The vertical distribution of APP (1 and 4 m deep), examined for 9 months at North Richmond, showed that overall mean density at 1 m was significantly higher than overall mean density at 4 m. Overall, the cell density of APP at 1 m deep was positively correlated with temperature and total chlorophyll a. The present results suggest that APP may need to be incorporated into a conceptual model of river plankton food webs. Manuscript received 8 August 2000, accepted for publication 22 November 2000. KEYWORDS: environmental factors, Hawkesbury-Nepean River, phycoerythrin-rich picocyanobacteria, plankton food webs. INTRODUCTION In fresh waters, the presence of autotrophic picoplankton (APP, cell size: 0.2-2.0 um, Sieburth et al. 1978) has been reported from lakes of various trophic states in the Northern and Southern Hemispheres (Paerl 1977; Stockner and Antia 1986; Burns and Stockner 1991; Weisse and Kenter 1991; Jasser 1997; V6r6s et al. 1998). The ubiquitous and often abundant presence (>10° cells ml!) of APP in lakes has prompted many ecological studies to investigate the relationships between the population dynamics of APP and physico- chemical and biological conditions (see Weisse 1993 for review). Some studied also have focused on the trophic role of APP in aquatic food webs, especially in the context of Proc. Linn. Soc. N.s.w., 122. 2000 80 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER other microzooplankton (Stockner 1991; Weisse 1993). In terms of the pattern of associations between environmental factors and APP in lakes, the abundance of APP may positively correlate with temperature (Pick and Carron 1987; Kennaway and Edwards 1989) but negatively correlate with intensity of zooplankton grazing (Weisse 1988; Fahnenstiel et al. 1991). The relationship between APP abundance and lake productivity may vary according to the trophic states (i.e. positive relationship in oligotrophic lakes and negative relationship in meso- and eutrophic lakes) (Stockner and Shortreed 1991: Burns and Stockner 1991). Thermal stratification may affect the vertical distribution of APP, largely confining their abundant presence above the thermocline (Fahnenstiel et al. 1991). In addition, these environmental factors may produce complex interaction effects on the temporal variation in the composition and abundance of APP (Rhew et al. 1999). Despite detailed ecological studies of lake APP, such studies are few for river APP. In the present study, basic ecological aspects of APP were examined for a year in the freshwater portion of the Hawkesbury-Nepean River, a regulated coastal river in New South Wales. The present study aimed to examine 1) the seasonal and horizontal (longitudinal) variation in cell density and cell type of APP, 2) the vertical distribution of APP, and 3) the pattern of seasonal associations between river environmental variables and cell densities of APP. MATERIALS AND METHODS Study Sites The Hawkesbury-Nepean River flows through the Illawara range to its mouth north of Sydney; the river has a catchment area of 32000 km? and a main channel length of about 320 km. The river flow has been regulated by five major dams on its headwaters and partly by more than 13 weirs. The present study was conducted at three freshwater sites. They were Penrith (non-tidal, about 180 m wide and 2 m deep), North Richmond (tidal limit, 120 m wide and 6 m deep) and Sackville (tidal, 200 m wide and 6 m deep) (see Fig. 1 in Kobayashi et al. 1998 for locations of sites). Sampling, Enumeration and Cell Type Measurement At each site, four replicates of water sample (100 ml each) were collected by using a Haney-type trap (Gawler and Chappuis 1987) from a depth of 1 m between 10:00 and 14:00. At North Richmond, additional samples were collected from a depth of 4 m between September 1992 and March 1993 to investigate the vertical distribution of APP (i.e. between 1 m and 4 m deep). All samples were immediately fixed with a 2% filtered (0.2 um pore size) buffered-formaldehyde solution (buffer: sodium tetraborate), and were transported to the laboratory with ice and stored at 4°C in the dark. In the laboratory, samples were initially filtered through a 3 um polycarbonate filter (25 mm in diameter; Millipore) to remove larger phytoplankton and zooplankton (Hawkey and Whitton 1991). Subsamples (3-10 ml) of these were then drawn onto a 0.2 um black polycarbonate filter (25 mm in diameter) under low vacuum pressure (<150 millibars). A cellulose acetate filter (pore size 0.45 um) was placed between the black polycarbonate filter and the filter holder as a backing to obtain an even vacuum (Hawkey and Whitton 1991). The black polycarbonate filter was then mounted on a glass slide, with a drop of immersion oil placed on top of the filter before gently affixing a cover slip. The APP cells (* 2 um in any dimension) were counted using fluorescence microscopy at a magnification of x1000, on a Nikon Diaphot-TMD inverted microscope, equipped with the standard G- 2A green excitation filter set (excitation filter EX510-560, dichroic mirror DM580 and barrier filter BA590) and a 100-W mercury lamp. The cells that fluoresced red were counted. These cells were assumed to be phycoerythrin-rich picocyanobacteria (MacIsaac and Stockner 1993). Thus, strictly speaking, the present study is most likely to have estimated a portion of the entire APP assemblage that may include eukaryotic cells. A minimum of 200 cells or all cells that appeared in a maximum of 40 fields of view were counted for Proc. Linn. Soc. nN.s.w., 122. 2000 T. KOBAYASHI, S. WILLIAMS AND A. KOTLASH ‘(QOOSTZ:T :2189S) sdew orydessodo} wo peos sem YIPIAA ‘[OUURYOS IOATI Jo yjdop pue yIpIM aJeutxo1ddy,, To=u CE=U 77=u €7=u '(GiGS-Gip) ‘(OZ €-r8'0) ‘(O'S8I-O'ST) ‘(6° L7-6 71) LEZ ESI S‘Or C61 Q pue 00Z I[EAyORS [z=u pc=u 77=u p7=u ‘(6°77Z-8'T) ‘(9S I-8€'0) ‘(0°6S-0'1Z) ‘(687-0 TT) puowyory 611 L60 Eve C61 g pue QZ YON €7=u €7=u GC=U €7=u ‘(7 b-€0) ‘(68'0-97'0) ‘(O'rL-0'S) ‘(¢°L7Z-0'8) 07 Lv'0 8°ZI 9°07 7 pue OST yyUuag (-l 31) (-l Sn) (uw) D “ll sw) snioydsoyd (9.) «4idop yAydoiojyD uaso3 [RIO], [210], aimesoduia |, pue YIpIAA aS SUONeAIISGO JO JoquINU dy} = U “UMOYS oe SasoyjUsed Ul odURI puke URI. “EG66[ ISNSNy pue Z661 Joquiajdas udaMJOq JOATY URIdaN-AINQsayMeH{ IU} UI SOTQUIIVA [PJUDWIUOIIAUD JOF SONSIVe}S AJeWIUNS “] 9IqGeL Proc. Linn. Soc. N.s.w., 122. 2000 82 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER expressed as the number of cells ml". Between December 1992 and August 1993, the proportional occurrences of morphological cell types of APP were investigated from 50 cells randomly selected from the pooled sample on each sampling date at each site. River Environmental Variables Temperature (°C) was measured with a Yeo-Kal Model 603 oxygen/temperature meter at 0.5 m depth. Water samples were collected for analysis of total phosphorus (ug | '), total nitrogen (mg I') and total chlorophyll a (ug 1"') in a Niskin-type bottle at 0.5 m depth. The samples were analysed by the methods described in Clesceri et al. (1989). Data on flow rate (10° 1 day") over Penrith weir were provided by AWT Environmental Science and Technology Division. Statistical Analysis A simple correlation analysis was used to detect any significant association between river environmental variables and APP cell densities at each site (a=0.05). All data except temperature were transformed by log,, to meet the assumptions of normality and homoscedasticity. All data analyses were made using the SAS computer programs (Anon. 1989). RESULTS River Environmental Conditions Between December 1992 and November 1993, flow rate over Penrith weir ranged from 2.3 x 10’to 4.1 x 10°1 day" (Fig. 1). Overall, temperature was in the range 8-28.9 °C. The means and ranges of concentrations of total phosphorus, total nitrogen and total chlorophyll a increased downstream (Table 1). 5000 4000 Flow rate (10° L day ~') Figure |. Flow rates over Penrith weir in the Hawkesbury-Nepean River. Proc. Linn. Soc. N.S.w., 122. 2000 83 T. KOBAYASHI, S. WILLIAMS AND A. KOTLASH +8C 0 snQ0'0- snV0'0 +«*9D 0 OUP snOb 0 ater OF encom snob O- alo sn LO'0- ***L9O- snot O- «x08 0 snc lL O- #*SSO snt0'0- snCC O +bV 0 snvlO pv |{AydoioyyD uasontu eo; snioydsoyd jej0], amjeloduia |, MOLY [121200 oTPAyoeVS puowyory YON yyuued aS "JUROTIUSIS JOU = SN ‘x L00'0>4 {xx 10'0>4>100°0 ‘xS0'0>4>10'0 :S[aAa] DoURdIFIUSIC “Q/-99 = U ‘SJUIDIJJ909 UONL]IIIOD [[BI9AO JOY “JWOIOGJOO9 UONE[ILOS Yyoeo OJ pZ-1Z = U “! Bo] Aq powoysueN 1am ainjesoduray }daoxe sonjea [TY “JOATY URodan-AinqsoymeH{ 24} ul UoyyURTdooId s1ydoNOe Jo AyIsUOp [[90 ULIUT PUL SofqeIIeA [eJUSWIUOITAUD UI9MIJOq SJUIIOIJJIOO UOHE[IMO a[duIIg °Z AQeL Proc. Linn. Soc. N.s.w., 122. 2000 84 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER & Penrith ang) E a 8 2 © oA = @® [=] oO 92S fe) N D 93J F M A M J J A North Richmond ae | E 2 8 =) P= (77) <= @ a 4” Sackville a 3 E 2 8 2 =) = 4 i— @® a oO 92S Oo N D 93J F M A M J J A Figure 2. APP in the Hawkesbury-Nepean River: Solid lines = mean density + SD at 1 m deep (n=4); dotted lines at North Richmond = mean density + SD at 4 m deep (n=4). Proc. Linn. Soc. n.s.w., 122. 2000 T. KOBAYASHI, S. WILLIAMS AND A. KOTLASH 85 APP at 1 m depth APP were present at all three study sites throughout the year (Fig. 2). Cell density tended to be high between mid-summer and early autumn at Penrith and North Richmond. Over the sampling period, cell density varied two orders of magnitude from 2.2 x 10° to 32) xallO7icells! male There were three morphologically distinctive cells: coccoid, ellipsoid and rod-shaped. In terms of proportional occurrences, coccoid cells became more important downstream from 25 to 52% of total cells during the study, whereas ellipsoid and rod-shaped cells decreased downstream from 64 to 47% and from 11 to 1% of total cells, respectively (Fig. 3) Vertical Distribution of APP between 1 and 4 m depth Between September 1992 and May 1993, the cell density of APP at 1m deep ranged from 1.5 x 10* to 3.2 x 10° cells ml! (overall mean 1.2 x 10° cells ml') and the cell density of APP at 4 m deep ranged from 8.4 x 10? to 2.7 x10° cells ml! (overall mean 9.4 x 10* cells ml") (Fig. 2). There was a significant positive correlation in cell densities of APP between depths (r=0.94, n=18, p<0.0001). Overall, the mean cell density of APP at | m deep was significantly different from that at 4 m deep (p=0.0055, n=18, paired-sample ¢ test [two tailed] on log, ,-transformed cell density data). Correlation between River Environmental Variables and APP Cell Densities The pattern of associations between APP cell density and river environmental variables differed between sites (Table 2). The strongest correlation was found between APP cell density and temperature at North Richmond. Overall, the APP cell density was weakly but significantly positively correlated with temperature and total chlorophyll a (Table 2). DISCUSSION The present study is the first to report the presence of abundant APP, in the freshwater portion of a regulated coastal river in Australia and elsewhere. The range of recorded cell densities of APP in the Hawkesbury-Nepean is within the range reported for lakes of the Northern Hemisphere (Stockner 1991; Weisse 1993; Szelag-Wasielewska 1997) and is comparable to that in lakes of New Zealand (Burns and Stockner 1991). A change of two orders of magnitude in seasonal cell densities of APP at study sites also is within the range reported for temperate lakes (especially mesotrophic-eutrophic) where APP abundance may change seasonally by almost four orders of magnitude (Weisse 1993). As has been reported for lakes (e.g. Fahnenstiel et al. 1991), the vertical heterogeneity of APP abundance exists in the Hawkesbury-Nepean River, at least at North Richmond. In lakes, higher densities of APP near the surface water are often observed during summer, coinciding with the development of summer thermal stratification (Stockner 1991; Gaedke and Weisse 1998). In the present study, the vertical heterogeneity of APP was examined only for limited duration at a single site. It is difficult to clearly demonstrate such a seasonal pattern in the vertical distribution of APP in the Hawkesbury-Nepean River. Nevertheless, the surface APP cell density tended to be higher than the deep APP cell density at North Richmond in January and February during the summer of 1992 and 1993 (Fig 2). The three morphologically distinctive types of APP cells are present in different proportions in time and space in the Hawkesbury-Nepean River. A variety of morphological cell types of prokaryotic APP (including coccoid, ovoid and rod-shaped) has been reported for lakes, but there is currently no key available for the identification of proper APP species (Maclsaac and Stockner 1993; Weisse 1993). This is chiefly because the majority of APP “forms” are prokaryotes that lack cell organelles and internal structures useful for Proc. Linn. Soc. N.s.w., 122. 2000 86 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER 100 Penrith 80 40 Cell density (%) 20 100 North a Richmond 40 Cell density (%) 20 100 | Sackville 80 Cell density (%) 20 Figure 3. Relative cell densities (%) of morphologically distinctive APP in the Hawkesbury-Nepean River. Horizontal shading = coccoid cells; vertical shading = ellipsoid cells; black = rod-shaped cells. Proc. Linn. Soc. n.s.w., 122. 2000 T. KOBAYASHI, S. WILLIAMS AND A. KOTLASH 87 the identification of species (Weisse 1993). Moreover, Weisse (1993) notes that even the morphologically uniform phycoerythrin-rich cyanobacteria may consist of more than one species. The taxonomic status of APP in the Hawkesbury-Nepean River awaits further studies. In the present study, there was no strong, consistent pattern of temporal associations between river environmental variables and cell densities of APP among study sites. The positive correlation of APP cell density with temperature at Penrith and North Richmond is, nevertheless, in accord with similar findings for some of the temperate lakes (Kennaway and Edwards 1989; Burns and Stockner 1991). Nutrients and total phytoplankton biomass (as measured by total chlorophyll a concentrations) also showed a certain degree of correlation with APP cell densities. On the other hand, river flow, which is often strongly associated with the seasonal variation in density of river microplankton (Kobayashi et al. 1998 and reference therein) was not a significant correlate of APP at any of the three study sites. Overall, the relatively low correlation coefficients indicate that large variability is associated with the seasonal relationship between the examined environmental variables and APP cell densities in the Hawkesbury-Nepean River. In an overview of APP from marine and fresh water ecosystems, Stockner (1988) has stressed the necessity of incorporating APP into conceptual models of lake plankton food webs, especially in the pelagic zone of ultra-oligotrophic systems. In these systems, APP may be key components in carbon metabolism and energy transfer, along with their heterotrophic counterparts (bacteria). Although this view is based on studies in freshwater lakes, APP may also need to be incorporated into a conceptual model of river plankton food webs. As in other rivers, microzooplankton predominate in the Hawkesbury-Nepean (Kobayashi et al. 1998). Many species of microzooplankton have been reported to effectively consume picophytoplankton (Stocker and Antia 1986; Weisse 1988; Miiller et al. 1991). Thus, APP may occupy an important trophic niche in plankton food webs of the river. In New Zealand lakes, Burns and Stockner (1991) have observed many picophytoplankton cells even in the gut of the small cladocerans such as Ceriodaphnia dubia and Bosmina meridionalis, although the digestion of the ingested cells by the cladocerans was not clearly demonstrated. Bosmina meridionalis occurs in the Hawkesbury-Nepean River and the microzooplankton community of the river consumes small algal food particles of ~5 um in diameter (Kobayashi et al. 1996). Further studies of the ecology of APP are warranted, especially in relation to the trophic role of APP in the plankton food webs in the Hawkesbury-Nepean River. ACKNOWLEDGEMENTS We thank Professor C. W. Burns, University of Otago for advice on microscopic observation of au- totrophic picoplankton. We thank Dr R. Oliver, Corporative Research Centre for Freshwater Ecology for comments. This work was partly supported by the Strategic Resources Planning Branch of Sydney Water Corporation. Proc. Linn. Soc. N.s.w., 122. 2000 88 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER REFERENCES Anon. (1989). ‘SAS/STAT User’s Guide’. Version 6, 4th edn. (SAS Institute: Cary, NC). Burns, C.W. and Stockner, J.G. (1991). Picoplankton in six New Zealand lakes: abundance in relation to season and trophic state. 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Seasonal changes in chlorophyll-containing picoplankton populations of ten lakes in Northern England. /nternationale Revue der gesamten Hydrobiologie 76, 545-554. Jasser, I. (1997). The dynamics and importance of picoplankton in shallow, dystrophic lake in comparison with surface waters of two deep lakes with contrasting trophic status. Hydrobiologia 342/343, 87-93. Kennaway, G.M.A. and Edwards, G. (1989). Seasonal and vertical distribution of picoplankton in Llyn Padarn, Gwynedd. British Phycological Journal 24, 375-384. Kobayashi, T., Gibbs, P., Dixon, P.I. and Shiel, R.J. (1996). Grazing by a river zooplankton community: importance of microzooplankton. Marine and Freshwater Research 47, 1025-1036. Kobayashi, T., Shiel, R.J., Gibbs, P. and Dixon, P.I. (1998) Freshwater zooplankton in the Hawkesbury-Nepean River: comparison of community structure with other rivers. Hydrobiologia 377, 133-145. Maclsaac, E.A. and Stockner, J.G. (1993). Enumeration of phototrophic picoplankton by autofluorescence microscopy. In ‘Handbook of Methods in Aquatic Ecology’, (Eds. P.F. Kemp, B.F. Sherr, E.B. Sherr and J.J. Cole). (Lewis Publishers: Boca Raton, Fl). Miiller, H., Schone, A., Pinto-Coelho, R.M., Schweizer, A. and Weisse, T. (1991). Seasonal succession of ciliates in Lake Constance. Microbial Ecology 21, 119-138. Paerl, H.W. (1977). Ultraphytoplankton biomass and production in some New Zeland lakes. New Zealand Journal of Marine and Freshwater Research 11, 297-305. Pick, FR. and Caron, D.A. (1987). Picoplankton and nanoplankton biomass in Lake Ontario: relative contribution of phototrophic and heterotrophic communities. Canandian Journal of Fisheries and Aquatic Sciences 44, 2164-2172. Rhew, K., Baca, R.M., Ochs, C.A. and Threlkeld, S.T. (1999). Interaction effects of fish, nutrients, mixing and sediments on autotrophic picoplankton and algal composition. Freshwater Biology 42, 99-109. Sieburth, J., Smetacek, V. and Lenz, J. (1978). Pelagic ecosystem structure: heterotrophic compartments of the plankton and their relationship to plankton size fractions. Limnology and Oceanography 23, 1256- 1263. Stockner, J.G. (1988). Phototrophic picoplankton: an overview from marine and freshwater ecosystems. Limnology and Oceanography 33, 765-775. Stockner, J.G. (1991). Autotrophic picoplankton in freshwater ecosystems: the view from the summit. Internationale Revue der gesamten Hydrobiologie 76, 483-492. Stockner, J.G. and Antia, N.J. (1986). Algal picoplankton from marine and freshwater ecosystems: a multidisciplinary perspective. Canadian Journal of Fisheries and Aquatic Sciences 43, 2472-2503. Stockner, J.G. and Shortreed, K.S. (1991). Autotrophic picoplankton: community composition, abundance and distribution across a gradient of oligotrophic British Columbia and Yukon Territory lakes. Internationale Revue der gesamten Hydrobiologie 76, 581-601. Szelag-Wasielewska, E. (1997). Picoplankton and other size groups of phytoplankton in various shallow lakes. Hydrobiologia 342/343, 79-85. Voros, L., Callieri, C., Balogh, K.V. and Bertoni, R. (1998). Freshwater picocyanobacteria along a trophic gradient and light quality range. Hydrobiologia 369/370, 117-125. Weisse, T. (1988). Dynamics of autotrophic picoplankton in Lake Constance. Journal of Plankton Research 10, 1179-1188. Weisse, T. (1993) Dynamics of autotrophic picoplankton in marine and freshwater ecosystems. Advances in Microbial Ecology 13, 327-370. Weisse, T. and Kenter, U. (1991). Ecological characteristics of autotrophic picoplankton in a prealpine lake. Internationale Revue der gesamten Hydrobiologie 76, 493-504. Proc. Linn. Soc. n.s.w., 122. 2000 Reproduction in the Short-beaked Echidna, Tachyglossus aculeatus: Field Observations at an Elevated Site in South-east Queensland Lyn A. BEARD AND GORDON C. GRIGG Department of Zoology and Entomology, The University of Queensland, Queensland 4072, Australia. Beard, L.A. and Grigg, G.C. (2000). Reproduction in the short-beaked echidna, Tachyglossus aculeatus: Field observations at an elevated site in south-east Queensland. Proceedings of the Linnean Society of New South Wales 122, 89-99. As part of a radiotelemetric study of echidnas (Tachyglossus aculeatus) in south-east Queensland focussing on thermal relations, we were able to confirm and extend present knowledge of echidna reproduction. Mating was concentrated in July and August, as elsewhere, but we found that echidnas have the ability to conceive successfully a second time within the one season, apparently in response to losing the first young. Echidnas in this area of south-east Queensland may be able to attempt breeding every year. Our data supports published estimates of gestation in the range of 20 to 23 days. Females spent two to three weeks in a plugged ‘incubation’ burrow, maintaining a high and stable body temperature for a period encompassing the last few days of gestation, all of incubation and the first few days of the hatchling’s life. The single young was carried in the female’s pouch for 45-50 days, attaining a body weight of approximately 200g before being stowed in a different plugged ‘nursery’ burrow. We describe the first detailed timing of a female’s visits to suckle her young. She visited regularly, every six days at first, gradually increasing in frequency to about every four days before the visits ceased and, presumably, the newly-independent young emerged at a calculated five and a half months of age. Manuscript received 16 August 2000, accepted for publication 22 November 2000. KEYWORDS: echidna, monotreme, radiotelemetry, reproduction, Tachyglossus aculeatus. . INTRODUCTION Short-beaked echidnas (Tachyglossus aculeatus) are extremely cryptic in the wild and rarely breed in captivity. This makes studies of their reproductive biology difficult and piecemeal. Early information was gained from dissection of dead specimens, single opportunistic observations on animals in the wild and sequential observations of females and their young taken into captivity. This identified the breeding season (July-August) and the egg incubation period (10.5 days) and provided a detailed picture of the anatomy and biochemistry of the reproductive organs and lactation (see Griffiths 1968, 1978). However, comprehensive information on life history parameters and behaviours was lacking. The advent of radiotelemetry made it possible to obtain information about reproduction by echidnas in their natural habitat. However, early telemetric studies concentrated on other aspects of echidna biology such as home range (Augee et al. 1975). Some short-term telemetric studies by Green et al. (1985) and Griffiths et al. (1988) Proc. Linn. Soc. N.s.w., 122. 2000 90 REPRODUCTION IN ECHIDNAS added information on milk intake by echidna offspring and suckling behaviour of lactating females, as did an observation on a single female echidna by Abensperg-Traun (1989). Long-term radiotelemetry studies of echidnas in the wild began in 1986, focussing on thermal relations (Grigg et al. 1989, 1992) and also providing some of the first longer- term field observations on echidna reproduction (Beard et al. 1992). Shortly thereafter, Rismiller, working on Kangaroo Island off the southern coast of Australia, commenced an extensive long-term study on echidnas in the wild, concentrating on reproduction and behaviour (Rismiller and Seymour 1991; Rismiller 1992, 1999; Rismiller and McKelvey 2000). Further fieldwork which may contribute to knowledge of echidna reproduction is currently underway also in Tasmania (S. Nicol and N. Andersen, University of Tasmania, pers. comm.) While a number of the questions about echidna breeding have now been answered, there is still uncertainty about many aspects, especially those dependant on direct and/or long term observation. Our continuing radiotelemetric study of echidna thermal relations in south-eastern Queensland has provided an opportunity to make such observations and to compare and contrast them with what is known of echidna reproduction and behaviour in more southern areas of Australia. MATERIALS AND METHODS The study site comprises parts of several grazing properties between Texas and Stanthorpe in SE Queensland at an elevation of 500-1000 metres, centred on 28°41’S, 151°32’E. The area is a mixture of mostly cleared, undulating grazing paddocks with varying grass cover depending on the season, and scrubby, mostly uncleared gullies. Echidnas were captured opportunistically and, with clearance from the University’s animal experimentation and ethics committee, were implanted with temperature-sensitive radio transmitters (Austec Enterprises, Canada and Sirtrack Ltd., New Zealand). The transmitters, fitted with an internal loop antenna, were coated in a biologically inert wax mixture (“Elvax’ / paraffin wax 20%/80% w/w) and implanted in the peritoneal cavity. They served to locate echidnas by radiotracking as well as to telemeter body temperature. Signals were acquired using a vehicle-mounted omni-directional whip antenna and TR-2 receiver + TS-1 scanner (Telonics, USA) and/or tracked on foot with a hand-held H- antenna (Telonics, USA) and receiver. One female also had, in two consecutive years, a waterproof, wax-coated temperature-sensitive datalogger (‘Tidbit’, Onset Corp., USA) implanted in the peritoneal cavity. Combined weight of the transmitter plus datalogger was approximately 50gm in an animal of average weight 3.25 kg. Body temperature data from animals with implanted transmitters could often be recorded automatically using a system which consisted of a timer switching on and off a receiver/scanner and tape recorder at preset intervals (Grigg et al. 1990). The same system could be used to monitor time spent by a female in a burrow, using a low gain antenna, or a feedline alone, placed on the ground above the burrow, so that it would pick up a signal from the female only when she was in the burrow or very close, during entry and exit. Any condition or activity which may have been related to breeding was noted for each animal during the course of tracking and weighing for other studies. As echidnas are normally solitary, aggregations of more than one animal were assumed to signal possible breeding-related activity. The presence of pouch young was obviously evidence of breeding, while an enlarged, loose pouch was taken to indicate its very recent vacation by a burrow-sized young (Griffiths 1968). Other signs included swollen mammary glands which were obvious when the female’s belly was exposed and from which droplets of milk could often be extruded, especially under anaesthetic for transmitter implantation. A swollen cloacal area in males was also taken to indicate some reproductive activity because, in this condition, the penis often partly everted when the animal was handled or Proc. Linn. Soc. n.s.w., 122. 2000 L.A. BEARD AND G.C. GRIGG 91 anaesthetised. Normally the penis, when not in use, sits in a fully-enclosed sac off the cloaca. We inferred dates of successful matings by working backwards from the estimation of the ages of pouch young using growth curves from Griffiths (1978, in litt.) and Green et al. (1985) and/or time in a brooding burrow, assuming an incubation period of 10.5 days (Griffiths 1978) and a gestation period of 20-23 days. We use the term gestation to describe that period between fertilisation and when the egg is laid. Fertilisation was assumed to occur shortly after mating (see Rismiller and McKelvey 2000). Pouch young were sometimes removed from the pouch for weighing, often with the female anaesthetised lightly with halothane until she relaxed enough for the young to be removed without a struggle, weighed and then replaced. The young were apparently insulated enough by the pouch to escape the effects of the anaesthetic. RESULTS Twenty-one echidnas, from a total of 30 animals (20 males and 10 females) which were captured in the study area over nine years, provided observations related to breeding. Individuals remained in the study population for lengths of time varying from a few months to several years. Those we lost presumably had transmitters fail or emigrated (Table 1). Table 1. Echidnas included in this study. Echidna # Sex | Captured Lost BW at capture | Time followed (kg) (months) 52 AES a atop 0 Pee Rs 26.4.90 19.1.91 4.45 IE a a FS ee eee dies Ocal eae Cee er OO aioe Seles week ac LSS kaos dutnetieds lee SS SSE a ee noo ae 2.8.90 19.12.90 3.8 4.5 23.10.92 11.10.93 4.0 IHS) 6.9.90 22.8.92 2.85 11.10.90 11.10.93 2295 36 19.6.91 11.10.93 28 0S i Ea Bea ne ee eee Sosa Doig ME nae F28ei oon e. ibm osintin Raseaiw an) i 4.3 (10.2.92) 22.9.92 29.10.92 20.7.94 20.7.94 9.7.95 | present _| 23.11.95 | present 30.11.95 present 19.3.96 present [mi [ 9110196 | present | ade tee Proc. Linn. Soc. N.s.w., 122. 2000 92 REPRODUCTION IN ECHIDNAS Mating and its timing. Mating activity in this area is focussed in the second half of July and in August (Fig. 1). However, we saw males keeping company with females for longer than this (Fig. 1), so the potential for mating extends outside this period, as in the case of a female which, in October, replaced an egg lost earlier (see below). This second mating must have occurred on or about 22 October, making it the latest mating yet recorded. If, as we have assumed, a swollen cloaca is a guide, males are available and ready both before and after the season in which most matings are focussed (Fig. 1). Most commonly, mating in this area occurred two to three weeks after emergence from hibernation. However, some females mated within a week of emergence, as was typical in the Kosciusko study (Beard et al. 1992). In one instance we observed a male and female (#52) together less than six days after the female had emerged from hibernation (Fig. 2). Abrasions noticed around the cloaca of the female after this encounter suggested that this was a mating and subsequent observations of the size of the young indicated that it was successful. We have never found more than two males with a female at the same time. Nor did we see evidence of the mating ruts or trenches described on Kangaroo Island (Rismiller and Seymour 1991). Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec >< >< ex Neob ae Gaeta 50% in Antarctica under spring “ozone hole” conditions (Madronich et al. 1998). This depleted ozone layer does usually not _extend as far north as Australia, but stratospheric winds can occasionally carry ozone- depleted air masses towards Australia causing a short term rise in UVB values. Although the relative rise in UVB has been most pronounced in the polar regions over the last decade (Kerr 1994), high ambient doses of UV-radiation are characteristic of tropical/ subtropical continents such as Australia even under normal stratospheric ozone concentrations (Fleischmann 1989). In these regions, the light path for solar radiation is short and the usually clear, oligotrophic water column exhibits a high transparency for UVB (Smith and Baker 1979). Consequently, many phototrophic organisms in aquatic ecosystems may be affected by this spectral waveband (Franklin and Forster 1997). Multiple harmful effects of UVB on marine primary producers have been reported, and include the direct influences on molecular targets such as nucleic acids and proteins, on physiological processes such as photosynthesis, growth and on community structures (Smith et al. 1992; Buma et al. 1995; Davidson et al. 1996; Franklin and Forster 1997; Aquilera et al. 1999). Of major interest is the identification of repair and/or protective mechanisms that allow phototrophic organisms living in high-light habitats to survive and reproduce. Proc. Linn. Soc. N.s.w., 122. 2000 124 MAAs IN RED MACROALGAE An important physiochemical mechanism against biologically harmful UV-radiation involves the biosynthesis and accumulation of photoprotective sunscreens. Typically absorbing in the UVA (320-400 nm) and UVB, these compounds were invoked to function as passive shielding substances by dissipating the absorbed radiation energy in form of harmless heat without generating photochemical reactions (Bandaranayake 1998). The most common substances with a potential role as UV-sunscreens in marine organisms are the mycosporine-like amino acids (MAAs), a suite of chemically closely related, water-soluble compounds. MAAs have been identified in taxonomically diverse marine organisms including bacteria, cyanobacteria, micro- and macroalgae, invertebrates and fish (Dunlap and Shick 1998). Their function as intracellular screening agents has been inferred from a decrease in concentration with increasing depth as observed in corals (Dunlap et al. 1986) and macroalgae (Karsten et al. 1999). In addition, macroalgae from South Europe contain up to 2-fold higher MAA contents compared to similar species from higher latitudes indicating a positive relationship with the natural solar UV-radiation of the respective biogeographic region, 1.e. the higher the UV-dose the more MAAs are formed and accumulated (Karsten et al. 1998a). In more recent studies on microalgae, Riegger and Robinson (1997) calculated sunscreen factors for Antarctic phytoplankton due to the presence of MAAs of up to 0.72, 1.e. 72% of harmful UV quanta were absorbed before hitting intracellular molecular targets. In the red-tide dinoflagellate Gymnodinium sanguineum Hirasaka, MAAs prevent, at least partially, UV-induced inhibition of photosynthesis (Neale et al. 1998). Although MAAs are widely present in various types of marine organisms, few data exist of their type and quantity in macroalgae (Nakamura et al. 1982; Karentz et al. 1991; Karsten et al. 1998a,b), in particular from high-radiation coasts such as in Australia. In the present investigation a qualitative and quantitative inventory was made of MAAs in red macroalgae collected from the rocky shore in southeastern New South Wales and southern Victoria. MATERIALS AND METHODS The locations of collection in southeastern New South Wales and southern Victoria are shown in Figure | and the red macroalgal species studied are listed in Table 1. All plants were sampled during a field trip in March 1999 directly from the shore as attached or drift material, or by snorkeling. Afterwards the algae were air-dried in the sun followed by storage in sealed plastic bags under cool, dry and dark conditions until analysis. Thalli of about 10-20 mg dry weight (DW) were extracted for 2 h in screw-capped centrifuge vials filled with 1 mL 25% aqueous methanol (v/v) and incubated in a waterbath at 45°C. After centrifugation at 5000 g for 5 min, 700 uL of the supernatants were evaporated to dryness under vacuum (Speed Vac Concentrator SVC 100H). Dried extracts were re-dissolved in 700 pL 100% methanol and vortexed for 30 s. After passing through a (0.2 um membrane filter, samples were analysed with a Waters HPLC system according to the method of Karsten et al. (1998a), modified as follows. MAAs were separated on a stainless-steel Phenomenex Sphereclone RP-8 column (5 um, 250 x 4 mm I.D.) protected with a RP-8 guard cartridge (20 x 4 mm I.D.). The mobile phase was 5% aqueous methanol (v/v) plus 0.1% acetic acid (v/v) in water, run isocratically at a flow rate of 0.7 ml min". MAAs were detected at 330 nm and absorption spectra (290-400 nm) were recorded each second directly on the HPLC-separated peaks. Identification was done by spectra, retention time and by co-chromatography with standards extracted from the marine red macroalgae Chondrus crispus Stackhouse (Karsten et al., 1998a) and Porphyra umbilicalis (Linnaeus) Kiitzing, as well as from ocular lenses of the coral trout Plectropomus leopardus (Lacepeéde, 1802), kindly sent by Dr. David Bellwood, James Cook University, Townsville, Australia. Quantification was made using the following molar extinction coefficients: shinorine: €334=44,700 (Tsujino et al. 1980), palythine: e320=36,200 (Takano et al. 1978), palythinol: €332=43,500 (Dunlap et al. 1986), porphyra-334: e334=43,300 (Takano et al. 1978), Proc. Linn. Soc. N.s.w., 122. 2000 U. KARSTEN 125 asterina-330: e330=43,500 (Gleason 1993). All amounts are given as mean of 4 replicates (+SD) based on separate extracts from separate algae, randomly collected from the respective habitat and expressed as concentration on a dry weight basis. New South Wales Victoria Melbourne Warrnambool Lonsdale Pacific Ocean 146° 150° FIGURE LEGENDS Figure 1. Map showing collecting location in southeastern New South Wales and southern Victoria, Australia. RESULTS The MAAs extracted from the dried red algal samples were characterised by HPLC, and identified and quantified according to their retention times, absorption spectra, co- chromatography with standards and molar extinction coefficients (see Materials and Methods). Five different MAAs could be detected within the samples investigated, all of which were identified as shinorine, porphyra-334, palythine, asterina-330 and palythinol (Table 1). The sum of all MAAs ranged in all macroalgae analysed from 0 (no trace) to 5.5 mg g! DW. While typical subtidal species such as Ballia callitrichia, Hypnea episcopalis, Nizymenia australis and Phacelocarpus alatus contained no MAAs at all or traces only, intertidal species such as Bangia atropurpurea, Capreolia implexa, Gelidium australe and Porphyra columbina exhibited high MAA concentrations between approximately 2.5 and 5.5 mg g! DW (Table 1). Quantitatively asterina-330 and palythinol played a minor role as indicated by low maximum concentrations of 0.38 mg g' DW as detected in Laurencia elata. While palythine showed high contents of up to 1.7 mg g'! DW in only few species such as L. elata, shinorine was the quantitatively dominant MAA in most species containing this compound. The maximum amounts of shinorine reached up to 3.9 mg g! DW. Porphyra-334 occurred in high concentrations between 1.5 and 2.5 mg g! DW in Bangia atropurpurea, Laurencia rigida and Porphyra columbina (Table 1). Proc. Linn. Soc. N.s.w., 122. 2000 MAAs IN RED MACROALGAE 126 Table 1 — Ultraviolet absorbing mycosporine-like amino acid (MAA) concentrations in red macroalgae collected in March 1999 from the rocky shores of southeastern New South Wales and southern Victoria. Values are given as mean + standard deviation (n=4) and expressed as mg per g dry weight; all MAAs are listed in terms of retention time. n.t.: no trace. EEE Species Collecting location Shinorine Porphyra-334 Palythine Asterina-330 __Palythinol XMAAs Amphiroa anceps (Lamarck) Decaisne Batemans Bay, NSW _—0..22+0.06 0.01+0.00 n.t. n.t. n.t. 0.2340.01 Amphiroa gracilis Harvey Warmambool, VIC 0.07+0,02 n.t. nt. n.t. nt. 0.07+0.02 Ballia callitricha (Agardh) Montagne Sorrento, VIC n.t. 0.02+0.01 n.t. n.t. n.t. 0.01+0.01 Ballia callitricha Port Lonsdale, VIC n.t. n.t. 0.01+0.00 n.t. N.t. 0.01+0.00 Bangia atropurpurea (Roth) C.Agardh Batemans Bay, NSW 0.11+0.01 2.54+0.29 0.03+0.01 n.t. n.t. 2.68+0.31 Capreolia implexa Guiry et Womersley Sorrento, VIC 2.36+0.36 0.06+0.03 0.79+0.13 0.14+0.02 n.t. 3.3640.52 Capreolia implexa Batemans Bay, NSW 3.85+0.71 0.04+0.01 1.33+0.13 0.24+0.03 0.04+0.01 5.48+0.73 Ceramium sp. Port Lonsdale, VIC 1.68+0.36 0.16+0.06 0.67+0.25 0.06+0.02 nt. 2.57+0.49 Champia sp. Warmambool, VIC 0.02+0.01 n.t. 0.02+0.01 0.01+0.00 n.t. 0.04+0.02 Cheilosporum sagittatum (J.V.Lamouroux) Port Lonsdale, VIC nut. n.t. n.t. n.t. n.t. n.t. Areschoug, Corallina officinalis L. Batemans Bay, NSW 1.03+0.23 0.01+0.00 0.09+0.03 0.01+0.00 n.t. 1.14+0.26 Dictymenia sp. Warrnambool, VIC 0.02+0.01 n.t. 0.01+0.00 n.t. n.t. 0.03+0.01 Gelidium australe J.Agardh Batemans Bay, NSW 2.15+0.02 0.01+0.00 0.01+0.00 0.01+0.00 n.t. 2.18+0.02 Gelidium australe Port Lonsdale, VIC 0.90+0.06 0.01+0.00 1.00+0.04 0.15+0.01 n.t. 2.05+0.03 Gelidium crinale (Turner) Gaillon Batemans Bay, NSW 2.92+0.23 0.02+0.01 1.05+0.03 0.18+0.01 n.t. 4.18+0.28 Gelidium pusillum (Stackhouse) Le Jolis Port Lonsdale, VIC 2.06+0.24 0.04+0.01 0.92+0.59 0.23+40.03 n.t. 3.25+0.80 Hymenema curdieana (Harvey) Kylin Warrnambool, VIC N.t. n.t. 0.01+0.00 n.t. n.t. 0.01+0.00 Hymenocladia chondricola (Sonder) Port Lonsdale, VIC n.t. n.t. n.t. n.t. n.t. n.t. Lewis Hypnea episcopalis Hooker et Harvey Warrnambool, VIC n.t. n.t. n.t. n.t. n.t. n.t. Jania micrarthrodia J. V.Lamouroux Port Lonsdale, VIC 0.98+0.07 0.01+0.00 1.15+0.14 0.06+0.01 n.t. 2.20+40.08 Jania sp. Sorrento, VIC nt. n.t. n.t. n.t. n.t. n.t. Laurencia botryoides (Tumer) Gaillon Sorrento, VIC 0.40+0.12 0.01+0.00 0.28+0.06 0.04+0.01 0.31+0.09 1.04+0.27 Laurencia elata (C. Agardh) Harvey Port Lonsdale, VIC 1.58+0.26 0.04+0.01 1.70£0.22 0.25+0.05 0.38+0.09 3.9540.38 Laurencia filiformis (C.Agardh) Montagne Port Lonsdale, VIC 0.34+0.05 0.01+0.00 0.64+0.03 0.05+0.01 n.t. 1.04+0.02 Laurencia rigida J. Agardh Batemans Bay, NSW 0.07+0.01 1.52+0.45 0.19+0.04 0.07+0.02 0.05+0.02 1,90+0.52 Laurencia tumida Saito et Womersley Port Lonsdale, VIC 0.54+0.22 0.01+0.00 0.63+0.22 0.04+0.01 n.t. 1.2340.45 Metagoniolithon stelliferum (Lamarck) Sorrento, VIC 0.68+0.08 0.02+0.01 n.t. 0.03+0.01 n.t. 0.70+0.08 Weber-van Bosse Nizymenia australis Sonder Port Lonsdale, VIC n.t. n.t. n.t. n.t. nt. n.t. Nizymenia conferta (Sonder) Port Lonsdale, VIC n.t. n.t. n.t. nt. n.t. nt. Chiovitti, Saunders & Kraft Phacelocarpus alatus Harvey Port Lonsdale, VIC n.t. n.t. nt. n.t. nat. nt. Plocamium angustum (J.Agardh) Port Lonsdale, VIC 0.25+0.11 0.01+0.00 0.36+0.22 0.02+0.01 n.t. 0.63+0.21 Hooker et Harvey Plocamium dilatatum J.Agardh Warrnambool, VIC 2.21+0.40 0.04+0.01 0.76+0.30 0.14+0.06 nt. 3.154072 Plocamium mertensii (Greville) Harvey Port Lonsdale, VIC 0.70+0.08 0.02+0.01 0.43+0.03 0.11+0.01 n.t. 1.26+0.11 Porphyra columbina Montagne Batemans Bay, NSW 0.92+0.13 1.88+0.35 0.16+0.05 nt. nt. 2,950.52 Pterocladia capillacea (S.G.Gmelin) Batemans Bay, NSW 2.40+0.40 0.04+0.01 0.05+00.01 0.04+00.02 n.t. 2.52+0.41 Santelices & Hommersand Rhodymenia australis (Sonder) Harvey Warmambool, VIC 0.08+0.05 n.t. 0.05+0.03 0.01+0.00 nt. 0.13+0.05 Wollastoniella sp. Warmambool, VIC nt. nt. nt. nt. nt. nt. Wrangelia velutina (Sonder) Harvey Warmambool, VIC 0.03+0.01 0.01+0.00 nt. nt. nt. 0.04+0.01 N.S.W., 122. 2000 LINN. Soc. Proc. U. KARSTEN 127 DISCUSSION This study provides the first comprehensive survey of the qualitative and quantitative occurrence of MAAs in red macroalgae from temperate Australia. In contrast to brown and green macroalgae, UV-absorbing substances have been widely observed in many species of the Rhodophyta (Sivalingam et al. 1974; Sivalingam and Nisizawa 1990; Wood 1989; Karentz et al. 1991; Maegawa et al. 1993; Molina and Montecino 1996; Karsten et al. 1998 a,b). In the present study, the MAA concentrations measured in typical intertidal algae such as Bangia atropurpurea and Capreolia implexa are approximately >20-fold higher compared to sublittoral species such as Ballia callitrichia. This is in good agreement with earlier reports on Rhodophyta from Arctic to warm-temperate localities (Maegawa et al. 1993; Karsten et al. 1998a) which indicate that species from deeper water exhibit only trace amounts or even lack these compounds. The red algae can tolerate a wider range of radiation levels than any other group of macroalgae. The deepest known plant is a coralline-like species found at 268 m off the Bahamas that grows at < 0.1 umol photons m* s (Littler et al. 1985). Other members of the group well reproduce and survive in the upper intertidal zone, often fully exposed to bright sunlight at >2200 umol photons m*® s! (Pedroche et al. 1995). Between these extremes the radiation quality and quantity reaching different species in different depths is highly variable due to the inherent optical properties of the water column, sun angle, latitude, season and weather conditions. However, sublittoral red algae are adapted to the generally low under-water radiation climate and hence are characterised as ‘shade-plants’ (Raven et al. 1979; Liining 1990). These species usually exhibit a lower photosynthetic capacity and rate of dark respiration than ‘sun-plants’, as well as optimum growth at low photon flux densities. Moreover, photosynthesis of macroalgae from deeper waters was shown to be particularly sensitive to UV radiation (Bischof et al. 1998). Since sublittoral plants are generally never exposed to high irradiances including UV, at least not for long periods, there is no physiological need to synthesise and accumulate metabolically expensive MAAs as indicated in the data presented. This in turn would save energy to better support other essential pathways such as, for example, light-harvesting phycobilisomes. It had been recently reported from Malaga in southern Spain (36.6°N - similar latitude as the locations in this study) that the depth distribution of brown macroalgae on the shore is controlled by the incident UV-radiation due to the species-specific sensitivity of spores against this short wavelengths (spores from shallow water species are more resistant than spores from species collected at greater depths). This means that one specific developmental stage of the life history is the main target of UV-radiation and this may affect zonation (Wiencke at al. 2000). Compared to sublittoral red algae, intertidal species are known to contain high contents of MAAs (Maegawa et al. 1993; Karsten et al. 1998a), which is in good agreement with the results shown. While most plants growing in this regularly exposed habitat are able to flexibly synthesise and accumulate these compounds in response to the respective radiation climate, some taxa such as Bangia atropurpurea exhibit always a high steady- state concentration. In this particular species cells seem to be loaded-up with the photoprotective substances, which is consistent with the typical occurrence very high on the shore. Besides the depth zonation, the biogeographic distribution of macroalgae seems to be another important factor controlling the MAA concentrations, since species from lower, high-solar latitudes always exhibit more MAAs than species from higher, low-solar latitudes (Karsten et al. 1998a). These observations indicate that the higher the natural solar UV-radiation of the respective habitat the more MAAs are formed and accumulated in these plants. MAAs are one of nature’s sunscreens, with 19 structurally distinct compounds so far identified in marine organisms (Dunlap and Shick 1998). Although MAA levels in Proc. Linn. Soc. N.s.w., 122. 2000 128 MAAs IN RED MACROALGAE macroalgae show a decline in concentration with increasing growth depth and are in general positively correlated with natural doses of UV-radiation (Karsten et al. 1998b), experimental evidence for the role of MAAs as UV-protectants in these plants is still circumstantial. Nevertheless, the presence of increasing MAA contents in the red alga Devaleraea ramentacea with decreasing depth strongly correlated with a more insensitive photosynthetic capacity under UV exposure (Karsten et al. 1999). Photosynthetic experiments on the unicellular microalgae Gymnodinium sanguineum proved that MAAs indeed act as spectrally specific UV-sunscreens (Neale et al. 1998). In marine invertebrates the function of MAAs as intracellular photon screening agents has been inferred from UV-induced delays in the first division of sea urchin embryos having low concentrations of MAAs compared to embryos with high MAA contents (Adams and Shick 1996). In another study, Dionisio-Sese et al. (1997) showed that the presence of MAAs in the surface tunic of the colonial ascidian Lissoclinum patella protect its photosynthetic symbiont, Prochloron sp., from UV-induced photodamage. Moreover, Ishikura et al. (1997) measured maximum MAA concentrations in the outermost surface layer of the siphonal mantle of the giant clam Tridacna crocea. The occurrence of MAAs in the animal tissue prevented an inhibition of photosynthesis of its zooxanthellae Symbiodinium sp., which outside the protecting animal tissue responded very sensitively to UV radiation. These authors calculated that the sunscreen capacity of the measured MAAs were sufficient to absorb 87% of 310-nm radiation and 90% of 320-nm radiation before reaching 0.2 mm depth in the siphonal mantle. All recent publications on marine algae and invertebrates strongly support the photobiological function of MAAs as a cellular defenCe system against the harmful effects of UV-radiation (Dunlap and Shick 1998). Therefore it is concluded that the physiological capability of intertidal red algae to synthesise and accumulate high MAA concentrations plays a vital role as biochemical adaptation ensuring survival under the environmental extremes in the habitat. ACKNOWLEDGEMENTS This project was financially supported by the Alexander von Humboldt Foundation via the Post-Contact- Programme and the Deutsche Forschungsgemeinschaft (Ka 899/3-2/3). The author likes to thank Linda Franklin, Heike Lippert, Alan Millar, Monica Schoenwaelder, John West and Joe Zuccarello for technical support, as well as for the identification of the species. REFERENCES Adams, N.L., and Shick, J.M. (1996). Mycosporine-like amino acids provide protection against ultraviolet radiation in eggs of the green sea urchin Strongylocentrotus droebachiensis. Photochemistry and Photobiology 64, 149-158. Aguilera, J., Karsten, U., Lippert, H., Philip, E., Vogele, B., Hanelt, D., and Wiencke, C. (1999). 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CMO%), Myo RORE AD see ties Eanes tayo Sahin Ree ame nwt auttioairs hie onlin won jo wurtsune bre adticloel (0487). f ,nwadedad brs. vidhaj Meret) 2. dtd BichS ian ened Venice laa ; , e avvniigiieboaies Yorigel ty, Luthar a A prameiMEAA. 2. wort ice Vuplererel So “ACE baw n sioes gamaskn snodtyynMndy - com wor freritailne sib 0 Ly SLT UL ECM, psa rien Pee ane Hak Bg ar ha ‘Aorta af ire? FI. 1d wen, & f. ayteent Mn ond oe ds Men 28 vee Naseal SL ees th deg of Aicdacrig phytgdandlen th tater’ aultate oe 1m i aie He a Ga eF OF vicar a. #8. biases. {als achi; § AS i fia ak + seme age ag gene ie debi, F } aie? MA. aa ant “ ews (GPR, saa ot aaem * gh Biogeography of the Freshwater Peracarida (Crustacea) from Barrington Tops, NSW Lorna T. ADLEM! AND BriAN V. Timms? '73 Bolwarra Park Drive, Bolwarra Heights NSW 2320 *School of Geosciences, University of Newcastle, Callaghan NSW 2308 Adlem, L.T. and Timms, B.V. (2000). Biogeography of the freshwater Peracarida (Crustacea) from Barrington Tops, NSW. Proceedings of the Linnean Society of New South Wales 122, 131-141. Distributions of certain groups of freshwater Peracarida (Crustacea: Isopoda; Amphipoda) in south-eastern Australia are known to favour high altitudes with associated cooler temperatures. Two species of crangonyctoid amphipod (Austrocrangonyx barringtonensis and A. hynesi) and two phreatoicid isopod species (Crenoicus harrisoni and Crenoicus n. sp.) have previously been documented from the Barrington Tops. During this study, six peracarid taxa were located including two new generic records for this area. These taxa showed interspecific variation in habitat and altitudinal preference on the Barrington Tops Plateau (~ 1585 m). The most influential environmental determinants of distribution for certain taxa were pH, flow rate and altitude according to canonical correspondence analysis (CCA). As a result, an altitudinally tiered distribution pattern could be seen on the plateau with Pseudomoera n. sp., the most tolerant taxon, occupying the widest range of altitudes and habitats. A broader investigation of peracarid distribution on the adjacent Nundle-Walcha Plateau to the north of Barrington and at Coolah Tops to the west, indicated the effects of past climate changes and remaining areas of refugia. Various levels of geographic speciation were identified relating to differences in adaptability and vagility between the amphipods and phreatoicid isopods. Manuscript received 29 December 1999, accepted for publication 22 November 2000. KEYWORDS : biogeography, climate change, peracarida, refugia, stenothermic, vagility. INTRODUCTION The Peracarida are of ancient lineage and are regarded as ‘living fossils’, with extant freshwater forms having undergone relatively little change from their ancestral marine relatives (Nicholls 1929). Two groups of present interest, the phreatoicid isopods and the crangonyctoid amphipods, have Gondwanan distributions and occur mainly in the cooler southern parts of Australia (Williams 1981, 1983). As might be expected from such distributions, many of the more northerly localities are at higher altitudes, especially among the amphipods. The Barrington Tops area (up to 1585 m asl) are known to have two amphipods, Austrocrangonyx barringtonensis and A. hynesi (Williams and Barnard 1988), and an isopod, Crenoicus harrisoni (Nicholls 1943). In addition, a species related to the recently described C. buntiae from the Boyd Plateau 300 km to the south-west (Wilson and Ho 1996) is known to occur at Barrington Tops (G. Wilson, pers. comm.). The aims of this work are: (a) to investigate the occurrences of Peracarida in the Barrington Tops and adjacent environs, map their distribution and assess their current status of abundance, and (b) to outline the habitat and altitude preferences of the taxa concerned. In addition, given the preference of many amphipods for cold temperatures Proc. Linn. Soc. N.s.w., 122. 2000 132 PERACARIDA FROM BARRINGTON TOPS and their use in environmental monitoring (Lake et al. 1979; Kangas and Geddes 1984), the mapping of present distributions may provide a basis for measuring the effects of future climatic change. STUDY AREA Barrington Tops Plateau (32°00’E, 151°30’S) is located 150 km north-west of the coastal city of Newcastle (Fig. 1). It is a remnant isolated paleoplain at 1000-1585 m asl and is bordered by steep escarpments (Pain 1983). The plateau surface is undulating with several, poorly drained valleys leading into deeply dissected ravines with steep stream gradients of the Manning River to the north and east and of the Hunter River to the south and west. Connected by a broad ridge, Gloucester Tops lies at 1313 m asl, while the disconnected Mount Royal at 1400 m asl represents the southern limit of higher altitudes. The northern high altitude limit is delineated by lower relief located at the Pigna Barney River which separates the Barrington Tops Plateau from a southern extension of the New England Plateau in the Nundle-Walcha district. This plateau has an altitudinal range of 1000 to 1400 m which cuts off steeply to the east and south with a gradual descent to the north, and is dissected by rivers and gorges. Drainage is into three major catchments; the Manning and Macleay Rivers to the east and south and the Namoi River to the west which constitutes part of the Murray-Darling system. A smaller isolated plateau (Coolah Tops 31°45’E, 151°05’S) lies 140 km to the west with an altitudinal range of 1000-1200 m and stream systems running into the Namoi and Macquarie Rivers (refer to Fig. 1). The Barrington Tops area has a cool-temperate climate with frequent frosts and occasional snow falls in winter. Rainfall is evenly distributed throughout the year and a strong precipitation gradient from 2000 mm p.a. occurs on the eastern side at Gloucester Tops to 1000 mm p.a. on the western side of Barrington Tops (Dodson 1987). The mean temperature range in July at 1300 m is -2.3 to 8.8°C with the January range at 9.1-22.8°C Sao e Walcha { @ Tamworth NSW Study Area x y) e@ Nundle UC es M4. 2m, @ Coolah S 9 @ Barrington Tops @ Gloucester » Gloucester Tops 32°05'S 4 State Forest Be National Park >z ) _ 20km Lae Maitland o— LS Ne e@ Cessnock wcastle ear 30'E Figure |. Location of the study area. Proc. Linn. Soc. N.s.w., 122. 2000 L.T. ADLEM AND B.V. TIMMS 133 (Dodson 1987). To the south, Mt Royal experiences an average rainfall of 1100 mm p.a., a mild to cool-temperate climate and temperature ranges of 15-30°C in summer and 3- 15°C in winter (Kinhill 1992). Barrington Tops supports a diverse range of vegetation from sub-alpine grasslands, open montane eucalypt forests, Sphagnum bogs and sedgelands on the plateau surface, to cool temperate rainforest and sclerophyll on the lower slopes. The plateau is used recreationally in National Park and State Forest areas with selective logging taking place in the latter. Private land in the northern area has been cleared for pastoral use. The majority of the Nundle-Walcha district is used for grazing with pockets of State Forest areas and pine plantations. Many swamps occur on private land, some of which have been drained to increase stock carrying capacity. METHODS A systematic search of the Barrington Tops and adjacent highland regions was conducted during the warmer seasons of spring and summer from February 1995 to March 1996. Sites were selected so as to include all major catchments, a range of altitudes and habitats, and any orogenic/ecological barriers within the areas which may affect population distributions. A total of 64 sites (alt. range of ~ 392-1500 m) were sampled in the Barrington Tops - Mount Royal area (Fig. 2). These sites were analysed to determine overall habitat preferences and distributional tendencies. A further 38 sites (alt. range of ~ 920-1345 m) from the Walcha-Nundle and Coolah Tops areas were investigated less intensively to provide information on distribution range and diversification of the peracarid taxa. Two different sampling techniques were used to collect amphipods and isopods. Sampling for amphipods employed a hand-held net with a mesh size of 1 mm positioned on the creek bed facing the current. A modification of “kick’ sampling (Frost et al. 1970; Chessman 1995) was used whereby rocks and underlying substrate were disturbed by hand causing the animals to be washed into the net. Contents were emptied onto a large white tray from which all specimens could be field picked. For isopods, a kitchen sieve with a mesh size of 1 mm was used for sampling along creek edges and in swamp areas. A “sieving and winnowing’ technique was used (G. Wilson, pers. comm.) that produced a clean sample from which to pick specimens. Sites yielding peracarid specimens were called ‘positive’ sites, while ‘negative’ sites gave no peracarideans. The collections (omitting actual sorting) were timed to assess abundance at each site. All specimens were transferred to vials containing 100% methylated spirits and stored for later identification and abundance counts. Ranked abundance was determined by dividing the number of specimens of each taxa in the sample by the time taken for the sample collection. These results were assigned to an abundance rank on a four point scale (see Table 2 and for details refer to Adlem 1996). At the Barrington sampling sites, temperature, pH, conductivity, dissolved oxygen and turbidity were measured using a calibrated Horiba water testing unit and momentary flow rate using a hydrometer. Degrees of exposure and turbulence were visually assessed and designated to a numerical scale (see Table 4). Overlying and underlying substrate were defined by diameter ranges, and vegetation (both aquatic and terrestrial) was identified and recorded at least to the genus level. Each immediate collection point was noted as a ‘midstream’, ‘edge’, ‘riffle’, ‘pool’ or ‘roots’ sample. Altitude and stream order were taken directly from 1:25 000 topographical maps as was distance from tributary sources using a curvimeter. Barrington site data were analysed using the FORTRAN program CANOCO for a direct correspondence analysis to indicate species/environment relationships. Canonical Correspondence Analysis (CCA) is a one-step analysis incorporating eigenvector ordination and multivariate direct gradient analysis (Ter Braak 1986). CANOCO (version 2.1) was Proc. Linn. Soc. N.s.w., 122. 2000 134 PERACARIDA FROM BARRINGTON TOPS % Pseudomoera (n.sp.) Crenoicus (nr) buntiae Austrocrangonyx spp. Crenoicus harrisoni a fe) ® Neoniphargus (n. sp.) 5 e Negative sites State Forest © National Park iB) National Park - proclaimed wilderness area Chichester State Forest Figure 2. Location of study area sites with respective Peracarida collected. run in a weighted centroid linear mode to combine physical data along which species data was distributed in accordance with influential environmental variables. Data such as exposure, temperature and dissolved oxygen were omitted from the analysis because of high variability owing to diurnal change. The resulting weighted mean scores were plotted on ordination axes to construct bi-plots. From these plots habitat preferences incorporating quantitative, ranked and qualitative data could be assessed and interpreted according to the positioning of corresponding coordinates. Proc. Linn. Soc. n.s.w., 122. 2000 L.T. ADLEM AND B.V. TIMMS 135 RESULTS Peracarida were collected from all major catchments. Four aquatic genera were found within the Barrington Tops study area, three of which were also present in the external exploratory areas in the Nundle-Walcha and Coolah districts. Terrestrial specimens were also found within collections (Table 1). Specimens with features approaching Crenoicus buntiae were collected from several sites and are referred to as C. n. sp. Both Austrocrangonyx hynesi and A. barringtonensis were present on the Barrington Tops, although a significant feature intergradation between the two species occurred with A. barringtonensis being predominantly identified (see Adlem 1996). Table 1. Taxonomic segregation of the peracarid fauna collected from the Barrington Tops Plateau. ; Aquatic/Terrestrial A/T A ORDER FAMILY GENUS SPECIES Amphipoda_ | Paramelitidae Austrocrangonyx | A. barringtonensis A. hynesi Eusiridae Pseudomoera n.sp Neoniphargidae Neoniphargus A. sylvaticus C. harrisoni n. sp. Talitridae Arcitalitris Isopoda Phreatoicidae Crenoicus Oniscidae (Tits Teeside] lary ar come eenal ? Neoniphargus sp. and Pseudomoera sp. in the Barrington region are new generic records and are of new species (J. Bradbury, pers. comm.). Sites sampled in the Nundle- Walcha district included further new species of Pseudomoera and Austrocrangonyx, and undescribed species of Crenoicus. Crenoicus spp. were the most frequent taxa encountered on the Nundle-Walcha plateau, being collected from 72% of positive sites. However, abundance ranks were generally low, particularly in swamp areas (see Appendix 2 in Adlem 1996). Pseudomoera sp. b was collected from a single site in this district which at present is the most northern locality for this genus. Crenoicus sp. was also found inland at Coolah Tops from a single site.but no amphipods were present in this area. Of the 64 sites sampled in the study area, 44 yielded collections of Peracarida. The sites in the Mount Royal State Forest yielded no specimens. Pseudomoera proved to be the most abundant (Table 2) and widespread taxon on the Barrington Tops plateau, being located at sites both within State Forest and National Park boundaries and externally on private land to the north. Austrocrangonyx, Crenoicus and Neoniphargus were all located at sites within these boundaries with Neoniphargus n. sp. and C. harrisoni (site13) co- occurring at the single locality in the upper reaches of the Manning River. Pseudomoera occurred at lower altitudes and consequently in higher order streams than the other taxa which were all found at distances <3 km from tributary sources at altitudes above 1000 m (see Adlem 1996). A broader distribution of Pseudomoera was also indicated by its occurrence within a wider altitudinal range (Table 2). The diversity of freshwater peracarid fauna increases with increasing altitude with the richest zone being 1400-1500 m. Therefore, 50% of the taxa found lie between 1200-1400 m. Austrocrangonyx is also relatively widespread and abundant (Table 2), but only occurred above 1100 m. Proc. Linn. Soc. Nn.s.w., 122. 2000 136 PERACARIDA FROM BARRINGTON TOPS TABLE 2. Number and percentage of positive sites (n = 44) occupied by peracarid taxa in the study area with abundance expressed as mean rank values. Mean abundance values ranked as 1 = lowest abundance; 4 = highest abundance. NA = not applicable (single sites). *Refer to Adlem, 1996. No. sites occupied /% | No. species exclusive Mean abundance rank sites / % value.* 2D Pseudomoera sp. 34 / 77.3% 24 / 54.5% A. barringtonensis 11 / 25.0% 1/ 2.3% C n. sp. 9 / 20.5% 8 / 18.2% A. hynesi 3 / 6.8% 0 Neoniphargus n. sp. 1/2.3% 0 C. harrisoni 1/2.3% The two bi-plots depicting species-environment relationships from the CANOCO analysis (Fig. 3) were produced simultaneously, therefore the respective plotted points represent integral ordination results. TABLE 3. Relationships between environmental vectors and ordination axes. 0.5129 0.6212 -) 0.0004 -) 0.1297 -) 0.4632 0.1741 Fig. 3a shows the plotted coordinates obtained for taxa and environmental vectors which are represented by lines. The increasing length of these lines indicates increasing magnitude of the respective vector, while position corresponds to the direction in which the vector is undergoing its greatest variability or change. The positioning among vectors relates to the degree of correlation between them. Therefore, vectors that oppose one another in the bi-plot have a negative correlation in regard to their respective increasing magnitudes. Hence, greater turbidities were negatively correlated with higher flow rates or more turbulent conditions, and less acid waters were negatively correlated with the higher altitudes on the plateau. Table 3 shows that the vectors with the longest lines (altitude, pH and mean flow rate) have higher correlation values with regard to the axes and are therefore the most influential factors towards the plotted positions of taxa and sites. Proc. Linn. Soc. n.s.w., 122. 2000 L.T. ADLEM AND B.V. TIMMS 137 A Axis 2 600 300 Mean Flow Rate Altitude eAustro Turbulence Axis 1 Turbidity eC. (nr) buntiae eNeo./.C.harrisoni 300 600 300 eAustro. ° ro} oie eC. (nr) buntiae eNeo./.C.harrisoni 300 600 Figure 3. Canonical correspondence analysis bi-plots showing the degree of correlation of species abundance with environmental vectors (A) and corresponding sites (B). Proc. Linn. Soc. N.s.w., 122. 2000 138 PERACARIDA FROM BARRINGTON TOPS Fig. 3b depicts plots of taxa in relation to corresponding positive sites where the respective taxa were located in greatest abundance. Altogether, Fig. 3 gives an indication of taxa plotting nearest to the vectors and sites to which they are most highly correlated, therefore the area of the bi-plot can be seen to represent a distance matrix originating from the mid-point, zero. Because all data are weighted linearly, the plots of Austrocrangonyx positive sites between altitude and mean flow rate show a correlation with higher altitudes, faster flow rates and greater turbulence. Opposing vectors to these sites are pH and conductivity, indicating lower values of these parameters (i.e. more acid, fresher waters) influencing the location of Austrocrangonyx sites. The most strongly corresponding site plot was sampled from roots of Sphagnum (site 21) with sites in shallow, stony riffle zones (21, 26a, 14 and 8) also distinctly conforming to abundant collections of this taxon. As shown in Fig. 3a, Pseudomoera plotting near the origin of the axes suggests all vectors as corresponding somewhat universally with this taxon, as the locus at the origin implies an equal influence of each vector, and no one vector is dominating the plot. Although plotting closely to conductivity and turbulence, these parameters did not correlate highly with either axis (Table 3), and therefore cannot be seen as strong influential factors in affecting Pseudomoera occurrences. Therefore, this central positioning reflects the ubiquitous range of Pseudomoera located with a greater variability of vectors implying a tolerance to a wide range of environmental conditions (excepting turbidity) in comparison with the other taxa as shown by raw data ranges shown in Table 4. Consequently, Pseudomoera was also found in a wide range of habitats which is supported by the majority of sites clustered around this taxon. These habitats ranged from deep, standing water within root systems of Myriophyllum aquaticum in muddy substrate (site 3), to fast- flowing, shallow riffle zones (2a and 38). However, the most abundant collection of Pseudomoera was at site 18 sampled from root systems of Nasturtium. Mossy rocks and liverworts in riffle zones of rainforest areas (24, 26, 27 and 45) also produced large numbers of this amphipod. TABLE 4. Raw data ranges of the environmental parameters analysed within which taxa were found in the study area. Turbulence values ranked as 1 = lowest degree of turbulence; 4 = highest degree. NR = not recorded. *Refer to Adlem, 1996. Parameter > Altitude Turbidity | Turbulence* | Conductivity Mean Flow (m) (NTU) (xS/cm) Rate Taxa | (cm/sec) Austrocrangonyx 1168-1490 11.3-42.1 Pseudomoera 865-1500 0.0-80.3 Neoniphargus 1400 i : NR C. n. sp 1260-1495 C. harrisoni 1440 The position of both Crenoicus harrisoni and Neoniphargus n. sp. coordinates plotted further away from vectors and sites. This is due to the rare occurrence of these species from a single site causing the plot to fall outside of the bounds of strong correlation and correspondence owing to a weaker data set. At site 13, Crenoicus harrisoni was observed to favour patches of gravel substrate within mud supporting Ranunculus and Montia, while Neoniphargus n. sp. was more prevalent among the root systems of these macrophytes. Proc. Linn. Soc. n.s.w., 122. 2000 L.T. ADLEM AND B.V. TIMMS 139 The plot of Crenoicus n. sp. at the end of the turbidity vector indicated the strongest correlation on the bi-plot. Strongly opposing vectors of mean flow rate and turbulence suggest a tendency of this taxa to prefer slow flowing or standing waters with greater turbidities (Table 4). This species of Crenoicus was collected from roots of Myriophyllum (sites 12 and 48), Ranunculus (site 39) and Sphagnum (sites 21a and 44). DISCUSSION This study has shown a greater diversity of freshwater Peracarida than has previously been recognised from the Barrington Tops Plateau, with at least six species located in the area. The occurrences of Austrocrangonyx spp. at higher altitudes above 1100 m in the uppermost headwaters demonstrates that Austrocrangonyx is a cold water, rheocolous taxon, favouring fast-flowing, shallow riffle zones, also noted in the Nundle-Walcha district during this study and by Boulton et al. (1995). The distribution of Austrocrangonyx was related to waters with lower pH values (Fig. 3a) which, at higher altitudes, are primarily influenced by the presence of the humic, peat-based swamps situated on the poorly drained plateau surface. Pseudomoera is the most commonly occurring and widespread taxon on the plateau, also occurring in the northern pastoral areas in creeks at lower altitudes with higher conductivities, greater turbidities, and higher pH levels associated with the absence of swampland and presence of exposed basalt surfaces. The high degree of abundance and occurrence of this amphipod would also be influenced by its ability to breed all year round as opposed to seasonal breeding in the other species (see Adlem 1996). The distribution of Crenoicus spp. on the Barrington Tops Plateau is strongly determined by the extent of Sphagnum areas and macrophyte establishment as populations were seen to favour these environs. Amphipods were also observed in the root systems of aquatic macrophytes common to the area (Myriophyllum and Ranunculus ) although not as abundantly as the phreatoicid isopods. Crenoicus sp. (nr.) buntiae preferred deeper, more turbid, slower flowing creeks with localised depositional areas of suitable substrate for the establishment of Myriophyllum aquaticum. Myriophyllum is known to occur prolifically in waters containing a high nitrogen content (Sainty and Jacobs 1981) which would become more available to the plants under acidic conditions (Salisbury and Ross 1985). Therefore, the acidic waters in the swamp vicinities contribute to a larger extent of available habitat for Crenoicus spp., from the fibrous peat and root systems of Sphagnum to basal sections of Myriophyllum in associated creeks draining out of and into swamp areas. Consequently, the most abundant occurrences of Crenoicus spp. were concentrated at sites located on the southerly plateau surface where swamps and alpine sedgeland communities dominate. Crenoicus spp. are widespread in the highlands of mid-eastern Australia (G. Wilson, pers. comm.), therefore their extensive occurrence within both the study area and exploratory areas is not surprising and new species on the adjacent plateau reflects strong divergence. The distributional ranges of certain taxa from the Barrington plateau proved to be much wider than formerly recorded. The discovery of a new species of Austrocrangonyx (J. Bradbury, pers. comm.) to the north on the adjacent Nundle-Walcha plateau indicates that this genus has a widespread, discontinuous distribution and is therefore not endemic to the Barrington area. Austrocrangonyx has also been found by one of us (BVT) near Ebor above 1200 m, which may well be the northernmost limit for this taxon. Freshwater amphipods have not been recorded from the Dorrigo State Forest area north of Ebor despite suitable altitudes (ca. 1380 m) (Chessman et al. 1994). Based on collection data for the Barrington-New England area however, the Nundle-Walcha plateau appears to be the northern limit of distribution for Pseudomoera. The catchment divides (which in these plateau areas are low) are unlikely to be Proc. Linn. Soc. N.s.w., 122. 2000 140 PERACARIDA FROM BARRINGTON TOPS barriers to dispersal for peracarids. The occurrence of Austrocrangonyx, Pseudomoera and Crenoicus spp. (e.g. Crenoicus n. sp.) in the Hunter and Manning catchments and also to a lesser degree in the Namoi which is part of the inland Murray-Darling system indicate a current widespread distribution. However, small scale isolation effects, particularly on the Nundle-Walcha plateau where agricultural and forestry operations are more prevalent, may contribute to habitat fragmentation and therefore loss of habitat area for peracarids. The fossorian Neoniphargus is likely to be a rare, relict species, and therefore an indicator of the Barrington area offering high altitude refugia for such forms. The cold water relict amphipod fauna mentioned by Williams and Barnard (1988) and Barnard and Barnard (1983) are included in the crangonyctid group of which both Austrocrangonyx and Neoniphargus are representatives. The occurrence of Crenoicus and absence of amphipods at Coolah Tops is significant. The only permanent waters are located at the head of the Talbragar River where Crenoicus was found. Though altitudes (and therefore temperatures) are marginal for amphipods (based on data from Barrington Tops) no amphipods, including the more tolerant Pseudomoera, were found. During a past arid phase, as for example ca. 18,000 years ago (De Deckker 1986), all streams in the Coolah Tops region would be intermittent at best and hence unsuitable for amphipods. Therefore, if amphipods ever existed at Coolah Tops, a past climate change would have caused their extinction there. However, phreatoicid isopods have the ability to sustain themselves at the sediment/groundwater interface, and could have survived dry periods. Lower altitudes, relatively dry conditions and temporary habitat also explains the lack of amphipods from the Mount Royal State Forest area. The upper reaches of tributaries where amphipods would be expected to be found were drought affected in this area. The distribution of the aquatic Peracarida observed in this study indicates that both Barrington Tops and the adjacent plateau in the Nundle-Walcha region maintain suitable refugia and habitat area for these crustaceans. The amphipod relict fauna appears to have a predisposition to colder environments and permanent waters which 1s corroborrated by their absence from Coolah Tops. Colder habitat and refuge areas on both plateaus are indicated by the widespread presence of Austrocrangonyx. The high altitude, cold water distribution of the amphipods may suggest that these animals could be monitors of possible future climate change, with Pseudomoera at the lower altitudes being a primary marker. The distribution pattern of the amphipods relates to colder temperatures in tributary headwaters and higher altitudes, whereas the phreatoicid isopods appear to have an overall wider habitat distribution and stronger divergence levels. The different distribution patterns between the amphipods and the phreatoicid isopods is related to their differing levels of vagility and adaptations to their respective environments. ACKNOWLEDGEMENTS Thanks go to Dr. John Bradbury for identification of specimens and also Dr. George (Buz) Wilson for identification and field techniques. Thanks also go to the numerous field assistants, the School of Geosciences at Newcastle University and the Linnean Society of New South Wales for supporting the study. National Parks and Wildlife Service and State Forests of NSW provided the research permits required and private landholders gave their permission to sample outside of these boundaries. REFERENCES ADLEM, L.T. 1996. Biogeography of the Freshwater Peracarida (Crustacea) from Barrington Tops, NSW. Unpublished Honours Thesis, University of Newcastle, NSW. BARNARD, J.L. and BARNARD, C.M. (1983). ‘Freshwater Amphipoda of the world. Vol. Il, Handbook and 3ibliography.’ (Hayfield Associates: Mt. Vernon, Virginia). Boulton, A.J., Kneipp, 1.J., Smith, A.P. and Sullivan, B.J. (1995). ‘Aquatic environment report Walcha/Nundle Styx River management areas. Walcha/Nundle and Styx River management areas EIS - Supporting document no. 3.’ (State Forests of NSW: Pennant Hills, NSW). Proc. Linn. Soc. n.s.w., 122. 2000 L.T. ADLEM AND B.V. TIMMS 141 Chessman, B., Growns, J., Hardwick, R., Holleley, D., Jackson, J. and Mcevoy, P. (1994). ‘Dorrigo three-year environmental impact statement area: Aquatic fauna report. Dorrigo interim EIS - Supporting document no. 2. Report no. 93/123.’ Compiled by Australian Water Technologies Science and Environment for State Forests of NSW. Chessman, B.C. (1995). Rapid assessment of rivers using macroinvertebrates. A procedure based on habitat- specific sampling, family level identification and a biotic index. Australian Journal of Ecology 20, 122-129. De Deckker, P. (1986). What happened to the Australian aquatic biota 18,000 years ago? In ‘Limnology in Australia’ (Eds P. De Deckker and W.D. Williams) pp. 487-496. CSIRO and Junk, Melbourne and Dordrecht. Dodson, J.R. (1987). Mire developments and environmental change, Barrington Tops, New South Wales, Australia. Quaternary Research 27, 73-81. Frost, S., Huni, A. and Kershaw, W.E. (1970). Evaluation of a kicking technique for sampling stream bottom fauna. Canadian Journal of Ecology 49, 167-173. Kangas, M.I. and Gedes, M.C. (1984). The effects of salinity on the distribution of amphipods in the Coorong, South Australia, in relation to their salinity tolerance. Transcripts of the Royal Society of South Australia. 108(3/4), 139-145. Kinhill Engineers Pty. Ltd. (1992). ‘Proposed forestry operations in the Mount Royal management area. Environmental Impact Statement. (Forestry Commission of NSW: Pennant Hills, Sydney). Lake, P.S., Swain, R. and Mills, B. (1979). Lethal and sublethal effects of cadmium in freshwater crustaceans. Australian Water Resources Council Technical Paper No. 37. Nicholls, G.E. (1929). Notes on freshwater Crustacea of Australia. Victorian Naturalist 45, 285-295. Nicholls, G.E. (1943). The Phreatoicidea Part I - The Amphisopidae. Papers on the Proceedings of the Royal Society of Tasmania 1943, 1-157. Pain, C.F. (1983). Geomorphology of the Barrington Tops area, New South Wales. Journal of the Geological Society of Australia 30, 187-194. Sainty, G.R. and Jacobs, S.W.L. (1981). “Waterplants of New South Wales.’ (Water Resources Commission: NSW). Salisbury, F.B and Ross, C.W. (1985). ‘Plant physiology.’ 3rd edn. (Wadsworth Publishing Company: California). Ter Braak, C.J.F. (1986). CANACO - a FORTRAN program for canonical community ordination by [partial] [detrended] [canonical] correlation analysis, principal components analysis and redundancy analysis (version 2.1). Technical Report: LWA-88-02 (Microcomputer Power: Ithaca, New York). Williams, W.D. and Barnard, J.L. (1988). The taxonomy of Crangonyctoid Amphipoda (Crustacea) from Australian fresh waters: foundation studies. Records of the Australian Museum Supplement 10, 1-179. Williams, W.D. (1981). The Crustacea of Australian inland waters. In ‘Ecological Biogeography of Australia.’ Vol 2 Part 4 - Inland Fresh Waters. (Dr. W. Junk bv Publishers: The Hague, Netherlands). Williams, W.D. (1983). ‘Life in inland waters.’ (Blackwell Scientific Publications: Melbourne). Wilson, G.D.F. and Ho, E.L. (1996). Crenoicus Nicholls. 1944 (Crustacea, Isopoda, Phreatoicidea) : Systematics and biology of a new species from New South Wales. Records of the Australian Museum 48, 7-32. Proc. Linn. Soc. N.s.w., 122. 2000 Ages, at ry: pa i = OF i ' he vaek cep ee sae: perenne MM (ira icicle a oe A wi a ads ; | ui Feguraeurat eee cen nnes fe el, Satie. dick ‘bap Ome” ee 5 peaenty OM bite eres atl 4 abel). ‘ail ' ie: Toye ORE rie ts cmt EH i Fr: he ak acnbuete we 4 e thie wea by et! wir } gh 4 wheeze ui vena ie meat GEAR i iif Hae eD Pasties himirds at nwa ai aldose RES cap ‘Wa a Nee iw aod si ig hale ic - xe WE - : fi fy oh : bie taycke xe! Aare li Asta nGwW ete i i Lan hes Nis sooty pemanene o ute rhe is ok rere Fe cet sa hyeratietl det ne ferro, esate Iie “hams iag ip ery | mia pout) jibe Fiat saan” yEee Fe cad BE Witt Wa Roh sae sk 4 x het pili We Ho saa ; 3s, Hu V Phone ate Ps i Picts 4 iia Ms Gaibinin walter ok hee erephiinnis Tite atyats We ef ew ihe ati i ries, NAP hog ens ihe pire avy’ eh erat " ae _ £ +9 Cle ai ; erin PASTTL iAARe sronrcr diveree a, E 1 Rhy he y ts eee rhe cw J iy ie Pike aiGicn ; if j 8 felt c d to teat MOLO TS ‘tor a : aA Pi yaa fi vae OCV Tet tite =fls, ie ys ANOWL pc EMENTS 7 elas! yur 4 tale geri Mai ti.oy spar ri itiela wetisd ae in corrsithaiy holt tox biden, Weehke ale gO the martes nis heh) a Aiaanin, Ihe Sanh og = weralty ex @ « Lannea Fouad of ow & ‘ Af Vghar’ Fins ays pr denge thet oipEY BN ' ; ce mad Slats reais ul SAY goto oeted Vie ‘r a yy perinlr i bolita ay a) yal ivr “haitle wor a ah ees -_ me “" ri inate (ealiae! = a rs al ed we PER! NCES : Pees Shan. Let oe Be Hh oak i rapa on ht eed Veruaide ae bia Garvin Rim cetera Fig er etre poeta news hie af Now ats, NE Beles = Hh RARS ‘ en Colt gh ery Tunah vance Adapgeti ot hr Wovild: wit ms . yy” decy edd Ametadaten BA), Verwon, Virgailia, Aap a Li." car bit wht Celie, BL) ealanG aivinioitrert > = Crammer nt oveae Walia tothe and Suva River namageiaiit we # 03 Vion of SOW, Pennant ithe, Wy, ¥ ey Notes on Successful Spawning and Recruitment of a Stocked Population of the Endangered Australian Freshwater Fish, Trout Cod, Maccullochella macquariensis (Cuvier) (Percichthyidae) J.W. DOUGLAS AND P. BRown. Marine and Freshwater Resources Institute, Snobs Creek, Private Bag 20, Alexandra VIC 3714, Douglas, J.P. and Brown, P. (2000). Notes on successful spawning and recruitment of a stocked population of the endangered Australian freshwater fish, trout cod, Maccullochella macquariensis (Cuvier) (Percichthyidae). Proceedings of the Linnean Society of New South Wales 122, 143-147. Conservation efforts for endangered fish species often include captive breeding programs that aim to re-establish viable populations in the wild. This study presents the first confirmed record of natural recruitment, to sub-adult, in a population of the endangered Australian freshwater fish, trout cod (Maccullochella macquariensis) derived from the stocking of captive- bred fingerlings. This represents a significant step in the conservation efforts for this species. Manuscript received 10 April 2000, accepted for publication 22 November 2000. KEYWORDS: conservation, captive breeding, endangered, fish, trout cod, Maccullochella macquariensis. INTRODUCTION Captive breeding programs that release progeny into the wild are common strategies used in the conservation of endangered fish species (Philippart 1995) The ultimate measure of success of such programs is the establishment of viable populations in the wild. Initial steps for this to be achieved include the development of breeding techniques, the placement of captive-bred animals where they can survive and grow, the initiation of natural spawning, and the recruitment and subsequent breeding in first generation individuals. Although not the only method for saving species, such programs are important conservation tools for fisheries managers, to be used in conjunction with habitat maintenance and protective legislation (Gooley 1992a). In Australia, artificial breeding techniques have been developed to assist conservation efforts for several threatened freshwater fish species, including the endangered trout cod, Maccullochella macquariensis Cuvier (Pisces: Percichthyidae) (Ingram and Rimmer 1992). BACKGROUND The trout cod is an endemic Australian fish considered threatened on an international level (Ingram and Douglas 1995). The species was once widespread throughout the southern Murray Darling River system of southeast Australia, but suffered a severe decline in range and abundance (Cadwallader and Gooley 1984). The species is restricted to only two isolated breeding populations (Ingram et al. 1990). Proc. Linn. Soc. N.s.w., 122. 2000 144 RECRUITMENT OF STOCKED TROUT COD National trout cod conservation efforts focus on protecting the existing populations with legislation and attempting to increase the number of self-sustaining populations through release of small fish produced from captive breeding programs (Douglas et al. 1994). Techniques to induce the species to breed in hatcheries were developed in the mid 1980s (Ingram and Rimmer 1992) and continued refinement of techniques has provided sufficient numbers of fingerlings, on a regular basis, to stock into selected waters. Since 1988 over 20 waters have been stocked with hatchery produced trout cod (Douglas et al. 1994). While there is evidence of liberated trout cod surviving to at least breeding age at many of the release sites (Douglas et al. 1994), there is no evidence of successful recruitment to adult from any of these populations. Preliminary evidence of spawning has been noted (Harris and Rowland 1996) from 1994 when a single larva (13.2 mm TL) and a single fingerling (92 mm TL) were sampled from two separate stocking sites in New South Wales. The larva was identified as a trout cod from diagnosis of myomere and pre-caudal vertebral counts (Brown and Neira 1998), and the wild origin of the fingerling was identified amongst fish of hatchery origin by the lack of enhanced otolith strontium concentration, which is used to mark hatchery-produced larvae in New South Wales (Brown and Harris 1995). Subsequent surveys of these two sites have found no further evidence of wild-bred juveniles or older year-classes. Loombah Weir (146°13’ 10” E, 36°43’ 18” S) was one of the original trout cod stocking sites in Victoria. Trout cod were not present in the weir prior to the stocking. Between 1988 and 1991, 8000 trout cod fingerlings, approximately 10-12 weeks old, were released into the only feeder stream above the impounded waters of the weir. Loombah Weir is a domestic water storage and was chosen because the catchment was relatively undisturbed and the area had limited public access. Non-destructive surveys between 1992 and 1995 monitored survival and growth of the stocked fish and recorded movement of fish downstream into the backed up waters of the weir. A monitoring survey in June 1998 using boat-mounted electrofishing produced a single adult (704 mm total length, > 5 kg) and two smaller trout cod (228 mm total length, 156 g and 199 mm total length, 91 g respectively) from the weir. The large fish was undoubtedly a survivor from one of the original stockings and was released. However, the size of the smaller fish implied they were likely to be younger than any of the previously liberated fish. Therefore both fish were sacrificed to estimate their ages from otolith sections. AGE DETERMINATION Age determination of the sampled fish was made by otolith reading and corroborated by length frequency analysis. Although the counting of annual growth increments in thin-sections of saggital otolith has not been validated for trout cod, it has been validated for the closely related species Maccullochella peelii peelii (Anderson et al. 1992a; Gooley 1992b) and Macquaria ambigua (Anderson et al. 1992b) and is likely to provide a valid method for estimating trout cod age. Thin sections of trout cod otoliths have previously been examined from over 70 juvenile and adult fish collected ad hoc from a variety of both natural and stocked populations. They show clear increments, which closely resemble those seen on the sibling species M. peelii peelii (S Morison, Central Ageing Facility, Queenscliff pers. comm.). Increment formation in M. peelii peelii occurs in September-November (Anderson et al. 1992a; Gooley 1992b). Examination of thin sectioned saggital otoliths from the two trout cod sampled from Loombah in 1998 revealed two opaque zones with a wide marginal increment (Fig. 1). This suggests that the fish were in their third year. The length-frequency distribution of trout cod, derived from a previous post-stocking survey in Loombah Weir in 1990 (Fig. 2) shows a size class between 210 and 300 mm total length. These fish were the oldest possible trout cod in the weir at the time and were in their third year (as trout cod had not been stocked into the site prior to 1988). The Proc. Linn. Soc. n.s.w., 122. 2000 145 J.W. DOUGLAS AND P. BROWN WU [ = Jeg aBOg “a8v JO SsIBaK € aq O} PAINS SI YSIJ OY], “ASPo OY) UO POULIOJ 9q 0} INOe SI JUDUSIOUT PITY & PUL SapIs [e.QUAA PUP [eSIOP YIOg UO IIqISTA Iv (e) SJUDUIDIOUI IvI[D OM, “3 G6 JYSIOM pure UW UWI 661 LL USIA ‘8661 ouNs ‘tam Yequioo’T Woy pafdutes poo jnoN kv JO YI]O}O [RISES B YSNOIY) UONSas asIOASULIL, | onst4 Proc. LInn. Soc. N.S.w., 1-2. 2000 146 RECRUITMENT OF STOCKED TROUT COD Figure 2. Comparison of length structure in samples of trout cod from a previous Loombah survey (1990) (clear bars) and the recently captured juvenile trout cod in 1998 (filled bars). Number of fish 12 N= 29 10 8 6 | A | | 0 + t t = t a + +——++} +++} 0 30 60 90 120 150 180 210 240 270 300 330 360 390 Total length (mm) lengths of the recently sampled trout cod (228 mm and 199 mm total length) are consistent with these 3-year-old fish, which provides additional confidence to the otolith based age estimates for these individuals. This age estimate suggests the birth year for the recently sampled trout cod as 1995, and from the known biology of trout cod, the date would be from October to November (Ingram and Rimmer 1992, Douglas et al. 1994) Since the stocking of trout cod in Loombah Weir ceased in 1991, the youngest possible age of stocked fish in 1998, in the impoundment, would have been seven years. Therefore, based on length and otoliths, the age estimates of the two trout cod sampled in 1998 indicate that these fish were younger than any stocked fish. Because no trout cod could gain access to the weir from other areas, the fish must be derived from a natural spawning of the previously stocked fish. This constitutes the first evidence of natural recruitment from a stocked population of hatchery-bred trout cod in Victoria and the first evidence of recruitment to three years of age from the natural spawning of any captive-bred trout cod population in Australia. DISCUSSION The time needed for the trout cod stocking program to produce viable breeding populations is unknown. However, because trout cod, like other large percichthyids, are likely to be relatively long-lived, the stocking programs should be viewed as long-term ventures. Loombah Weir was one the first waters stocked in the Victorian trout cod stocking program and it took nearly 10 years to observe some success. Monitoring of several other trout cod stocking sites nation-wide has also returned evidence of initial survival and growth of the stocked fish (Douglas et al. 1994) so it is likely that other sites may also show evidence of breeding and recruitment in the next few years. Proc. Linn. Soc. n.s.w., 122. 2000 J.W. DOUGLAS AND P. BROWN 147 Spawning and recruitment to three years of age is a positive step towards the aim of creating viable wild trout cod populations from releases of captive-bred, fish. Future monitoring in Loombah Weir should follow the progress of the naturally spawned generation towards this goal. ACKNOWLEDGMENTS We wish to thank Corey Green and staff at the Central Aging Facility for their prompt service in providing the otolith-based age estimates, and Sandy Morison, Lachlan McKinnon and Dr K. P. Sivakumaran who provided constructive advice on the manuscript. REFERENCES Anderson, J.R, Morison, A.K. and Ray, D.J. (1992a). Age and growth of Murray cod, Maccullochella peelii (Perciformes: Percichthyidae), in the lower Murray-Darling Basin, Australia, from thin sectioned otoliths. In: ‘Age Determination and Growth in Fish and Other Aquatic Animals’ (Ed D.C. Smith) Australian Journal of Marine and Freshwater Research 43, 111-142. Anderson, J.R, A.K. Morison and D.J. Ray 1992b. Validation of the use of thin-sectioned otoliths for determining the age and growth of golden perch, Macquaria ambigua (Perciformes: Percichthyidae), in the lower Murray-Darling Basin, Australia. In “Age Determination and Growth in Fish and Other Aquatic Animals’ (Ed. D.C. Smith) Australian Journal of Marine and Freshwater Research 43, 231-256. Brown, P. and Harris, J. (1995). Strontium Batch-Marking of Golden Perch (Macquaria ambigua) (Richardson) and Trout Cod (Maccullochella macquariensis)(Cuvier). In “Recent Developments in Fish Otolith Research’ (Eds D.H. Secor, J.M. Dean and S.E. Campana). University of South Carolina Press. Belle W. Baruch Library in Marine Science Number 19, 693-703 Brown, P. and Neira F.J. (1998). Family Percichthyidae. Basses, Perches and Cods. In ‘Larvae of Temperate Australian Fishes. Laboratory Guide for Larval Fish Identification’ (Eds F.J. Neira, A.G. Miskiewicz and T. Trnski,) pp. 259-265. (University of Western Australia Press: Perth). Cadwallader, P.L. and Gooley G.J. (1984). Past and present distribution and translocations of Murray cod Maccullochella peelii and trout cod M. macquariensis (Pisces: Percichthyidae) in Victoria. Proceedings of the Royal Society of Victoria 96, 33-43. Douglas, J.W., Gooley, G.J. and Ingram B.A. (1994). “Trout cod, Maccullochella macquariensis (Cuvier) (Percichthyidae). Resource Handbook and Research and Recovery Plan’. (Department of Conservation and Natural resources: Melbourne). Gooley, G.J. (1992a). Native fish stocking programs-What are the requirements? In ‘Proceedings of Symposium Freshwater Fisheries in Victoria- Today and Tomorrow’ (Ed. P. Cadwallader) (Department of Conservation and Natural Resources: Melbourne). Gooley, G.J. (1992b). Validation of the use of otoliths to determine the age and growth of Murray cod, Maccullochella peelii (Mitchell) (Percichthyidae), in Lake Charlegrark, western Victoria. In ‘Age Determination and Growth in Fish and Other Aquatic Animals’ (Ed. D.C. Smith) Australian Journal of Marine and Freshwater Research 43, 219-230. Harris, J.H and Rowland, S.J. (1996). Family Percichthyidae, Australian freshwater cods and basses. In ‘Freshwater Fishes of South-eastern Australia’ (Ed. R. McDowall) (Reed Books). Ingram, B.A., Barlow, C.G., Burchmore, J.J., Gooley, G.J., Rowland, S.J. and Sanger, A.C. (1990). Threatened native freshwater fishes in Australia- some case histories. Journal of Fish Biology 37 (Supplement A), 175-182. Ingram, B.A. and Douglas, J.W. (1995). Threatened fishes of the world: Maccullochella macquariensis (Cuvier, 1829) (Percichthyidae). Environmental Biology of Fishes 43, 38. Ingram, B.A. and Rimmer, M.A. (1992). Induced breeding and larval rearing of the endangered Australian freshwater fish trout cod, Maccullochella macquariensis (Cuvier) (Percichthyidae). Aquaculture and Fisheries Management 24, 7-17. Philippart, J.C. (1995). Is captive breeding an effective solution for the preservation of endemic species? Biological Conservation 72, 281-295. Proc. Linn. Soc. N.s.w., 122. 2000 eit itgtiabexhpeaegay tnienterslon ipa une bo? juOw bliw sidsit bonwage qisnsiag od to eeon79 5 ‘a a 6 ‘hbiueds VW dsdmaot at eh Reto. o Sea hia ss ac " pears — f sal i nh oc ee fee N28" | | i ATE OIA. 7s in ‘wobaren deyerverney Tiel + rae ge gniaes lenis any ‘eta bis Nand giao. hecities ieee risniurratevi2 TAK ba shacsragel lit staat nia? Lins watiistsizas: og8 beaks sions, a Bia tacuied i no oombs ovina 23 ae | . | ee axnvamss A ea Cole, 70 MWeTS Dike OB ts | yest bos AA cotoe Ae goon furery were : Hes MiB - 2 a rast ae, Coubuyttt sag count gftiniotigee rol athiio§ Taepl wh eh thebp ytartoisn B ‘aberhith Kp A ares) 23 eS ese Papen nes ts a a Yupagent tombaack A) Cesidins aeeN Ns) MIT aed loO to im HOTS Ht ni eimsarqgolovah wma? HE eho ps Aya (aito® as S| nasi. ae dngne ® ieaseinys Aermquire 5 2 Sig a = eae ON -ERO ek adele 2 a ate ara { Sate ums! Te 6 Vn. fat bor bas ee eee oueeeteiti Rd dee Las) jie’ nn wl oheuhe ta asada be bk Nati egdiaey! Riese gee peegipodinhons ae ip “ees aee air te at SE sang adnate —s sgt Aa tie, 5) Jomrameend), ay a Ia Us, MEO Tee Me ereecce a ce Neferani ‘nhgimat atten sermcieptenintite ae ati rapels. 12. nweecy Pre i ronatano ryt eieterary nae eh sirens his - oe Senaeens wae cies ai 55 lee te Sopa eoy ; certae aes wietny OTe IMC VIOUSLY? Sic ke Tee, PES. RIS oe Ws ul stg bartny ghar}, Sata ns aed eased) eRlaWhD DE cones ? BeOOR ((laeegy', hal) at a ne UNOS to 7% Vebutine HONEY un. ye 2h ollie Chad Velie ok. auto bie sy ane 1A bidet i ZEEE ay HaseAAN SARA WR SoA? vblicte RH? wetter Ce i hvu Dyer ee ileal yt, (bise a Onl! We eoitelt boa f AePe 1) WL, isigootl bh K Ag HE 2D eval) ous ion saitoonmmtieed .f oubiyehitalapeel) (QhR tort UA, i cumariabons mitt Ye Ginga jeu Han gap ow beoubal (2201) A.M rare fe . AAT: Won qushoonph.. vebieiaitsien: A) Gail siete Rane saath jodi: as milo hh ity, bondi tel Teale Alaa eso, insebi> 17 Prep, am! iu gat i nas As ieee likaly to he eeinuvery iam digo, ieritok ag PORT RINS sin b shine ay lng ‘=ilul a ewohudr Weg «ie omedhet fied woleiry Site Cha i the Wictorianl routed @ . a Arty a) of (ook tery 1) Veer Wo OCT omer BL CC RS, Mauniioni dy: of WEV . ie GA) cere dce4. cies apes tees, nishy ‘torte 4 evig nowt Aneel SUrVLY: bah sien aw! (Ar eitekeed fish Modulat eta (9oe7 ay Ht Ji Sikuly: that ier Atey mm pon ry ‘ tyecding and (acstenett in Ie Gast tour yearn, NOL 5h). od 2 oe Presidential Address for 1999-2000 Geodiversity: “green” geology in action R.A.L. OSBORNE School of Professional Studies, Faculty of Education, A35 University of Sydney, New South Wales 2006 Osborne, R.A.L., (2000) Geodiversity: “green” geology in action. Proceedings of the Linnean Society of New South Wales, 122, 149-173. Geodiversity is the whole range of natural Earth features and processes. Geoheritage consists of all the significant Earth features and continuing processes that we wish to keep, sustain, conserve, manage and interpret for their natural heritage value. The geodiversity practitioner is involved in all the phases of the geoheritage process: identification, documentation, conservation, management and interpretation. Identification can proceed by a variety of means, but is incomplete without field checking. Documentation not only involves describing the place, but also determining its significance. Determining significance is quite difficult, but can be aided and made more reliable by the use of systems of criteria. Description also entails determining the boundary of a place, using cadastral, natural, topographic, significance, catchment and natural system perimeters. Protective buffer zones may also be required. Conservation can be undertaken by legal means or by negotiation, but will not succeed unless there is management that produces continuous protective care of the significance of the place. Many attempts at legal protection have failed due to the lack of proper management. Interpretation is not only vital to increase public understanding of geoheritage places; it is an essential part of the conservation and management process. Geoheritage is a challenging area in which to work, requiring a broad knowledge of the Earth sciences coupled with expertise in, and commitment to, natural heritage conservation. Manuscript received 18 October 2000, accepted for publication 22 November 2000. KEYWORDS: environmental geology, geodiversity, geoheritage, geological heritage, geological monuments, natural heritage. Presidential Address for 1999-2000. Presented at the Royal Botanic Gardens, Sydney, 22 March 2000. INTRODUCTION When I meet people in Akubra hats and tell them that I am a geologist, they always ask me to give them a share of the gold I find. People in white construction helmets think I have come to solve their foundation or groundwater problems. Such are the popular images of geology and its role in society. Biologists, on the other hand, do exciting and important “green” things like saving whales, furry things and rare plants. There is however, a “green” branch of the Earth sciences variously called geodiversity, geoheritage, geological heritage, or Earth science conservation, concerned with saving the geological equivalents of whales, furry things and rare plants. Conserving, managing and interpreting significant Earth features is well advanced in the UK, USA and in former Eastern Block countries such as Slovenia and the Czech Proc. Linn. Soc. N.s.w., 122. 2000 150 PRESIDENTIAL ADDRESS 2000 Republic, but not in Australia. Here it runs a very poor third after biodiversity and cultural heritage. In 1996, the Australian Natural Heritage Charter established geodiversity as an essential element of natural significance to be considered in heritage conservation. This has had some impact at the Federal and Local Government level, but is yet to have a significant effect in New South Wales at the State level, where planning laws and land management policies are made. Geoheritage practice involves identifying places with potential significance, determining their extent and characteristics, assessing their significance and developing conservation, management and interpretation strategies. Most heritage workers lack the necessary Earth science background and most geologists and geomorphologists lack an understanding of the principles and practice of heritage conservation. WHY GEODIVERSITY? Sharples (1993 and 1995) introduced geodiversity into the Australian literature. Discussion by Dixon (1996), Wilkins and Osborne (1996) and Semeniuk (1997) followed. Geodiversity as a term has not been universally popular and during the 1990s its use caused a major split among geoheritage workers in Australia. Supporters of geodiversity were concerned that traditional approaches, as implied by terms such as geological heritage, were too narrow. Public and academic perceptions had greatly narrowed the range of features considered geological, often to the exclusion of important features such as landforms and soils. It was felt that a new term was necessary that encompassed the whole range of natural Earth features. The term, Abiotic, favoured by some conservation agencies, was also considered inappropriate as many Earth processes have a biological component. The Australian Natural Heritage Charter (Cairnes 1996) defined geodiversity as “the range of earth features including geological, geomorphological, palaeontological, soil, hydrological and atmospheric features, systems and earth processes”. Geodiversity is not intended to be a scientific concept. It is a technical term used in natural heritage conservation. Geodiversity does not imply that heritage conservation should particularly emphasise those places with the greatest range of Earth features. Geodiversity means identifying and conserving significant examples from the whole range of rocks, minerals, fossils, structures, landforms, soils, rivers, lakes, springs, etc., and places where Earth processes are occurring. Taken together biodiversity and geodiversity encompass the focus of this Society, “natural history in all its branches”, called the “whole realm of nature” by 18th century naturalists and hymn writers. PRACTICING GEODIVERSITY In New South Wales, and most other jurisdictions in Australia, geodiversity elements are not legally required to be considered in environmental impact statements, plans of management or state of the environment reports. While flora, fauna and archaeological surveys will be undertaken if a major development is proposed, geoheritage surveys are unlikely to occur. Most work for geodiversity practitioners comes from the public sector, particularly from Local Government and the Australian Heritage Commission. The work required is usually site specific, generating a few days work here and there, certainly not sufficient to make a living. Large jobs, which are rare, inevitably involve hiring casual staff; however finding people with a suitable background is not easy. Staff must understand local/regional geology Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 151 obvious choice, but today many have field areas in other countries, distant parts of Australia, on the seabed or under the ice. Often they have little knowledge or interest in the local or regional environment. Most have been taught not to read the local literature or papers more than five years old. It is possible to find suitable staff after considerable searching. The work of geodiversity practitioners is surprisingly similar to that of exploration geologists. The initial step of identification is exploration, but the object is not high- grade ore, rather places of significance. Determining the significance, condition and exact boundaries of the identified place is akin to finding the grade and tonnage of an ore body. IDENTIFICATION The first step is to determine what is significant and where it is located. While the step itself is obvious, how it should be done is not. A number of approaches have been taken each of which produces a particular type of outcome. Expert Polling Expert polling is a process by which experts in a field are asked to nominate places they consider significant to a list or sit around in a group and develop a list. This process is biased by those who choose to reply to requests for nominations or by those who are chosen to participate in workshops. Expert polling is a rapid and cheap way to produce lists of potential places for later investigation. This method was used by Percival (1979) to add 100 extra potential sites to the list of 100 geological sites previously assembled by the N.S.W. Geological Sites and Monuments Sub Committee of the Geological Society of Australia. Places identified by expert polling tend to have irregular spatial distribution (close to participant’s institutions or field areas) and low type diversity. Places west of Dubbo and in the New England region were poorly represented in Percival’s list, while Early Palaeozoic fossil localities and central volcanoes were predominant. Public Nomination Members of the public will often nominate places to the Register of the National Estate or council heritage registers. Some of these places are well known and recognised by the scientific community, while other places will have their significance substantially overstated. Sometimes local community members will nominate places that have not been previously recognised. Good examples are the Elizabeth Street Faults exposed in a road cutting in suburban Newport, north of Sydney. When residents brought the place to the attention of Pittwater Council in 1999, they thought the dipping sandstone beds were an outstanding example of cross bedding. Site inspection (Osborne and Osborne 2000) revealed that the beds were dragged down by a pair of normal faults not previously recorded either in the literature or on geological maps. Desktop Survey A desktop survey can be used where a large area is to be covered and funds are scarce. The fundamental assumption behind a desktop survey is that reliable and useful information can be found from the literature, maps, remote sensing, databases and other sources that can be brought to the desk. Desktop surveys produce lists of potentially significant places. Without field investigation, it is impossible to be sure of the existence, location, significance, condition or boundary of a place. An extreme example of a desktop geoheritage survey was undertaken as part of the Comprehensive Regional Forest Assessment process. The survey (Osborne et al. 1998) was of the Upper North East, Lower North East, South and Eden Regional Forest Proc. Linn. Soc. N.s.w., 122. 2000 152 PRESIDENTIAL ADDRESS 2000 Assessment Regions in New South Wales. The survey area covered most of the east coast and the eastern portion of the highlands of New South Wales, some 160,000 square kilometres, represented on ninety four 1:100 000 scale topographic maps. Over a period of four months the project identified 1,746 places of potential significance of which 1,241 (71%) had not been identified in previous surveys. Four months was insufficient to effectively cover all of the available literature and at least another six months would have been required to complete the project. Regional Approach Regional approaches are often favoured because they fit in with practical demands for planning information. The regional approach to geodiversity was considered so significant that the Australian Heritage Commission held a workshop on the topic in 1996 and the papers from it were published (Eberhard 1997). The success of the regional approach depends on how the regions are selected and defined, and on understanding the pitfalls inherent in the methodology (Osborne 1997). Regions based on catchments or local government areas may appear to be of little use for studies with a focus on bedrock geology, but such regions may be very important as they form the basis of land management. If regions are based on natural zonation (e.g. geological provinces) or given natural boundaries, it is absolutely essential to ensure that the significance of features located on the boundary is not ignored. Major faults and unconformities at the boundaries of geological provinces may be more significant than the rocks on either side of them. It would not be of much use if regional studies of the Sydney Basin and the Lachlan Fold Belt ignored the unconformity at Kanangra Walls, or if studies of islands and coastal areas stopped at the high water mark. Thematic Approach Thematic surveys, such as a survey of vertebrate fossil sites (Willis 1993) or my work on New England Karst (Osborne 1998), have the advantage that places are being identified and assessed by a specialist in the relevant area of study. The topics of thematic studies often reflect the availability and enthusiasm for conservation of specialists in particular fields, rather than any planned approach or decision about which themes need investigation. ASSESSMENT AND DOCUMENTATION Literature Survey Once a potential place has been identified it needs to be documented and assessed. The first stop is usually at the library to find out what, if anything, is known about the place. Looking backwards. Information about places of geoheritage significance is sometimes found on the World-Wide Web or in the latest journals. In most instances, however, the work of 19th and early 20th century geologists and naturalists needs to be consulted, often in rather yellowed volumes of this Journal. Other important sources include the Annual Report of the Department of Mines, Records of the N.S.W. Geological Survey and unpublished reports such as the Department of Mineral Resources GS series. On occasion the trail will lead to the dome of the Department of Lands building in Bridge Street, Sydney, where old maps and plans are stored. Often the historic literature will provide not only the best description and maps of the place, but also photographs from which the condition and integrity of quite small features can be judged. Edgeworth David’s work on glendonites at Huskisson (David et al. 1905) includes a detailed site map of the locality, which can still be used. Surprisingly some large boulders shown on his map continue to be useful reference points. The Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 11)3) photographs in David et al. (1905) allowed the subsequent survival of the glendonites in the rock platform to be evaluated. There appeared to have been little change or obvious deliberate damage between 1905 and 1996 (Osborne 1996). Reading between the lines The older literature is a great source of information about unusual and spectacular features. Writers in the older literature frequently commented on features that were not the prime focus of their research and described them in great detail even if they did not know what they were. Due to poor base maps and a tendency of some people to get lost, the location data is sometimes difficult to interpret and reading between the lines, tracing paths and finding out about non-current locality names is required. While most modern scientific writers know where they are, they don’t record much about anything that does not fit into their particular, very specialised, view of the world. A different type of reading between the lines is required here. Questions such as what soils or landforms might be associated with a particular rock type regularly need to be asked. Thank God for library angels Some places just don’t want to be found, and the literature doesn’t help. Several visits to the reported position of the Ramstation Creek limestone locality, near Dungog, between 1995 and 1998 failed to find any limestone. The map reference given on the relevant geological sheet (Roberts et al. 1991) seemed to match the location given by Jaquet (1901) and Carne and Jones (1919), but no limestone could be found. Just when I was about to give up and assume this was another nonexistent locality a library angel came to my rescue. A map (Jaquet and Harper 1899) fell out of a back pocket in Memoirs of the Geological Survey of New South Wales volume 2. The copy I had looked at previously had no map. The map not only showed the location and shape of the Ramstation deposit, about | km west of where I was looking, but also the location of three other deposits that had eluded me. It also became clear that although the Ramstation deposit had been described or noted by Carne and Jones (1919), Anon (1948), Lishmund et al (1986) and Roberts et al. (1991) none of the authors since Jaquet (1909) had actually been there and unfortunately neither have I. The super secret People love to have secrets. This is particularly the case with “special” places like fossil and mineral localities and limestone caves. The specimen or photograph seems to gain extra significance if “I can’t tell where it came from, but isn’t it wonderful”. Restricted circulation publications, strict membership criteria, secret maps and hidden databases are all used to restrict secrets to the few and “worthy”. Most secrets are known to a much wider population than their keepers ever imagine. Accessing “secret” information 1s rarely a problem, but deciding what to do with the information can be. Well-known places with no literature Many well-known features, both geological and geomorphological, are not mentioned in the scientific literature. Places nominated by academic experts often include their favourite student excursion localities. These are usually outstanding examples of some particular type of feature, but no one has ever bothered to describe them in a refereed journal. Tourist promoters, land managers and the public at large vote for iconic places with their lookouts, feet and cameras. What the public and the tourist industry consider important, however, is often quite different from what professional scientists value and describe. Some of the most visited places in New South Wales include the sea cliffs at North Head and the Three Sisters in the Blue Mountains. To my knowledge there is no published scientific literature on these features. As a consequence it can become quite difficult to demonstrate the significance of places that everyone agrees are significant. Location Finding the place If a location is mentioned in the literature it should be possible to pinpoint it on a Proc. Linn. Soc. N.s.w., 122. 2000 154 PRESIDENTIAL ADDRESS 2000 map and find it in the field. Published locations, however, are frequently wrong. Some reported occurrences simply don’t exist and some are duplicate records of other places, but with wrong locations. Most incorrect locations result from cumulative errors, poor initial reporting, mirror-image map copies, changing systems of grid references and poor or no archiving of data. Some of the most difficult problems arise when authors of compilations and review documents allocate precise locations to vague references given in original texts, without making any attempt to confirm the information. One team of compilers gave a precise grid reference based on statement in Carne and Jones (1919) that: “S.R. Beatty, District Surveyor, Maitland, has reported the occurrence of two deposits of limestone on the northern side of Arundle River, one about 9 and the other 10'/, miles W.N.W. of Copeland”. Another trap for the unwary comes from 1:100, 000 scale geological maps and their accompanying guidebooks. Most provide excellent information and location data. Some of these maps, however, extend over more than one standard 1:100, 000 sheet, and as a result over a grid zone boundary. In these cases the grid references on one part of the map (and in the notes) will not correspond to those on the standard 1:100, 000 and 1:25, 000 topographic maps for the same area. As a result of these and other difficulties, my survey of karst in the eastern New England (Osborne 1998) was not able to locate 15 out of 61 (25%) published limestone localities. Unrealistic expectations Land management authorities frequently have quite unrealistic expectations of what can be achieved from a desk survey. At best, a desk survey will give positions with an error circle of approximately 1 km ona 1:100, 000 scale map. That is assuming the place really exists. Those who can’t or won’t fund fieldwork often expect that desk surveys will not only produce precise grid reference data (+/- 10 m or 100 m), but also legal boundaries and management recommendations. These expectations are clearly a dangerous fiction. Ownership and management It is important to know who owns and who manages the places you wish to conserve. While it is fairly easy to determine who owns places with freehold title, increasingly state laws and local government planing instruments have a great influence on what you can do in your own back yard. It is vital to know not just who owns a place, but what the owners are legally allowed to do with it. Who actually owns and manages land in public ownership and land with less than freehold title is not always easy to determine. Frequently there are overlapping levels of management and disjunctures between legal precision and practical reality. It may be more important to discover who mows the grass and who empties the garbage bins than to know the name on the title, who pays the rates or which body holds the land in trust. It is essential to do the administrative searches and to talk to the person driving the tractor. One must never assume that fences, roads or even buildings are in the right place, that people really own their back yard or that land which the council manages as a park is a public reserve or council-owned land. Professor T.W.E. David unveiled a large painted wooden sign at Seaham Quarry, north of Raymond Terrace in 1926, which concludes; “Science trusts that the People of Seaham will kindly preserve this quarry intact for the benefit of future generations.” Everyone assumed that the quarry was public land, but surveys in the 1980s revealed that it was private property. The quarry was eventually purchased and is now part of Seaham Nature Reserve. The need for red lines on maps The world of land tenure and land management depends on red lines on maps. To conserve or manage a place requires a well-defined boundary that can be marked on plans and laid out on the ground by a surveyor. Locations defined entirely by a single grid reference, a dot or unbounded shading on a map will not do. While exploration geologists have great experience in pegging out claims, surprisingly some academic geologists and Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 155 geographers appear not to appreciate the importance of defining a place as an area (or volume) with a definite boundary. Small places and the problem of many maps It is often necessary to use a series of maps with differing scales to usefully locate small places. Fossil and mineral localities, some structures and springs are often less than a hectare, and may have an area of only a few square metres. While it may be possible to use a single plan to legally define their location, their significance will often relate to their regional or even continental geological or geomorphological context. As a consequence more maps may be required for the proper documentation of a small place than for a large one. Tenure blind or not? One of the most controversial issues in heritage identification is where one should look. Should places of significance be identified wherever they occur, or should land with some types of ownership or use not be evaluated for heritage significance? Some landowners, and categories of land users, argue the initial decision that land can be used for a particular purpose (residential, agricultural, forestry, mining) precludes it from subsequent heritage assessment. I, and many others, respond that heritage assessment should be tenure blind, particularly since many decisions about land use were made a considerable time ago, without any assessment or consideration of the impact of the designated use. This issue is particularly important in the case of land uses such as mining and waste disposal, where the designated use is likely to occur for a very brief period of time relative to the likely natural life span of either ecosystems or geoheritage features. Surprisingly, some state conservation agencies have argued that their reserves contain a complete and sufficient sample of all features of natural heritage significance in their state and that there is nothing of significance outside their reserves. Description A useful description must tell the reader what is there, allow them to recognise the significant features and understand why these features are important. Thinking about the audience Reports about places with geoheritage significance are rarely read, or used, by Earth scientists. They are mainly used by land managers, landowners and by council planning officers. Most of these people are unfamiliar not just with the language and concepts of the Earth sciences, but also with the idea that Earth features could be significant or worthy of conservation and management. Because professional conservationists, land managers and planners are so familiar with protecting and managing the living environment and the “rich tapestry of our priceless cultural heritage”, the description must highlight geoheritage significance in an unambiguous way. Object lessons of management not understanding what 1s significant at geoheritage sites abound. Examples include a landcare group planting trees on a naturally bare scoria cone, and millions of tourists visiting the lookouts at North Head being told about the shrubs behind them, but not about the cliffs or the view they went to see. Since the audience of the report is unlikely to recognise geoheritage features by name, (What’s a brachiopod, glendonite, fault, ria...?), maps, diagrams and photographs with scale, are an essential component of any description. The Statement of Significance The statement of significance is a key component of heritage listings, conservation plans and management plans. It is a concise statement about why the place is significant, and should form the basis for future conservation and management. The statement of significance must be technically precise, yet comprehensible to non-specialists. Proc. Linn. Soc. N.s.w., 122. 2000 156 PRESIDENTIAL ADDRESS 2000 A special style Since statements of significance have to be brief and contain a large amount of information, a particular style of writing has developed. The general form of these documents, usually less than an A4 page in length, is something like this: “The X (place) is an outstanding example of a Y (feature). It exhibits Z (rare or unusual characteristic) to a degree not seen elsewhere in the region. The place is largely undisturbed and sub feature | and sub feature 2 are found in a rare state of preservation....” For geoheritage places this style presents considerable problems. The readers are likely to have a reasonable understanding of statements like “contains species x and y, listed as endangered in New South Wales” or “ is the most intact surviving Victorian cemetery”. They are far less likely to understand or value “one of the few examples of Tertiary leucitite in Australia’. Writing a statement of significance forces you to consider why a feature is significant and then to explain this concisely. Condition and Integrity Condition It is important for a report to describe the present condition of a place. Is it a pristine forest, is ita mass of noxious weeds or an abandoned quarry partly filled with metallic farm waste and old bottles? It is important for the report to focus on the condition of the significant features at the place. The significance and condition of geoheritage features is usually unaffected by impenetrable noxious weeds, which often protect rather than harm. As a consequence what might be a disaster zone to an ecologist, may be a site in excellent condition to a geodiversity practitioner. Integrity It is important to distinguish between condition and integrity, as both factors my have a bearing on the significance of a feature. A single fossil of the whole organism although in poor condition may be more significant than a large deposit of well-preserved pieces (e.g. a whole trilobite vs lots of pygidia, an intact crinoid vs thousands of columnals). Integrity becomes an important issue if a significant place is modified or damaged after it has been documented and placed on a heritage register. How much can the integrity of a place become compromised before it looses its significance? This difficult question can only properly be answered if the condition and integrity of the place were well documented initially. Current condition vs threat It may be clear that there are threats to the condition and integrity of a place. While some make efforts to evaluate threats, others consider that documentation should only consider the place’s current condition and integrity. There have been two responses to dealing with places that are clearly at risk. The usual response is to say that if a significant place is threatened, then there is a strong case for documentation, listing and protection. The less common response is to do nothing where places are likely to be compromised or destroyed by a known legal activity, because it has already been decided that they will be destroyed. Boundaries One of the most difficult and important issues is where to draw the boundary. In conservation, planning and land management the position of a boundary has important legal and financial implications. Heritage listing or changes in zoning may be positive or negative to landholders’ interests in the order of millions of dollars. This makes it very important to determine a boundary that not only will result in the place being conserved, but can also be defended before administrative tribunals and the courts. A confusing outcome of different approaches being taken to boundary definition is that some places have multiple entries with different boundaries in the Register of the Proc. LINN. Soc. N.s.w., 122. 2000 R.A.L. OSBORNE 157 National Estate. One geological example is the Warrumbungle Volcano in central New South Wales. The Warrumbungle National Park is listed on the register and defined by its cadastral boundary. The Geological Society of Australia’s nomination of the Warrumbungle Geological Site, also listed, is based on a boundary designed to include all significant features following Percival (1979). This is a much more complex boundary, and covers a larger area than the national park, including areas of freehold land outside the park boundary. A number of different approaches can be taken when defining a boundary, each of which has quite different consequences for conservation and for people with an interest in the affected land. Cadastral boundary The simplest method of defining a boundary is to follow land tenure boundaries. If most, or a significant part, of a feature is in a reserve, national park, road reserve or within a single freehold Portion or Allotment, then the boundary off the title plan becomes the boundary of the place. This approach has two real advantages; the boundaries are already legally defined and only one landowner has to be dealt with. The disadvantages of using cadastral boundaries can be considerable. Significant natural features, particularly landforms and geological structures, are rarely restricted to a single rectilinear Lot or Portion. Similarly, processes that are likely to impact on the conservation of a feature are not often restricted to its exact physical location. Topographic boundary Topographic features such as streams, cliff lines and ridge tops would appear to make good boundaries, but where do you actually draw the line? Should the boundary be the top of the cliff, the base of the cliff, or some distance out from the base of the cliff so as to include rockfall and scree? While these types of boundaries are easy to plot from air photos and topographic maps, they are not so easy for surveyors to measure and define in the field. Boundaries based on contours are likewise attractive, but imagine constructing a boundary fence along a contour. Inclusive significance boundary If our aim is to “retain the natural significance of a place” (Cairnes 1996, p 10), surely it makes sense to draw a boundary that includes all its significant elements, irrespective of topography and land ownership. Inclusive boundaries are easy to justify, but often have complex and inconvenient shapes. These boundaries take no account of the surrounding environment or of practical issues such as tenure and management. Inclusive boundaries can result in “shrink wrapping”, which produces small discrete sites whose context is not retained. These are extremely difficult places to manage. Where a feature is unrelated to its surrounding environment, is very small, or is an isolated remnant, “shrink wrapping” is the only practical alternative. A classic example is the Dalton Fossil Leaf Deposit (Percival 1985), which consists of a single boulder of fossil- bearing rock, housed in a wire cage beside the local tennis court in the village of Dalton, southern New South Wales. Exclusive significance boundary An exclusive boundary is produced by looking at a large defined area in which significant features are distributed and then drawing a boundary that excludes those parts of the area which lack significance. Exclusive boundaries will frequently produce a pattern with patches of land with no significance surrounded by, or embayed into, significant areas. Exclusive boundaries can be useful tools for planning development within areas of generally recognised significance, such as national parks or heritage precincts. Buffer zones Buffer zones are areas that should be managed in order to conserve the significant places that they adjoin or surround. Buffer zones may be needed to control erosion, protect catchment areas or to provide a physical barrier against people, machinery or vehicles. Since buffer zones generally lack significance themselves, their creation needs to be Proc. Linn. Soc. N.s.w., 122. 2000 158 PRESIDENTIAL ADDRESS 2000 carefully justified. Natural system (ecological) boundary When ecosystems are being documented for conservation purposes it is normal practice to define boundaries that include, or attempt to include, the whole of the ecosystem in the area of identified significance. Such an approach can be taken with some Earth features, particularly active landform systems. The Earth system boundary of a beach could be drawn to include back dunes on the landward side and sand reservoirs in banks some distance out to sea. System boundaries of a river would include its catchment and estuary, and a karst by its catchment, sink and resurgence. Even if for practical and political reasons a place cannot be formally bounded by its natural system boundary it is useful for buffer zone management to define a natural system boundary. The adjacent place problem Many related natural features are not directly adjacent to each other, but separated by land with quite different characteristics. If related features are tens of kilometres apart, it makes sense to consider them as separate places for conservation and management purposes. If, however, related places are a few kilometres or less apart, practical and administrative issues can arise. Should adjacent, related features be considered elements of the same place, or should they each be considered to be a separate place? While state governments have been prepared to declare national parks and proclaim reserves composed of numerous disconnected parcels of land, the Australian Heritage Commission and others who keep heritage registers have often found dealing with related disjunct elements a difficulty. Most heritage registers were designed to deal with buildings with a discrete location and street address, not features such as chains of volcanic hills, or even small patches of remnant rainforest on the north coast of New South Wales, where this problem initially arose. Since the Register of the National Estate lists “places”, and gives them grid references, latitudes and longitudes, how, the bureaucrats ask, can a place have more than one location? SIGNIFICANCE What do we mean by significance, and how can we measure or determine it? Joyce (1995), a geodiversity sceptic, considered that “the significance of a geological feature or site lies in its value in research, reference or education at the local, national, international or world level.” This definition relates only to utilitarian scientific and educational values. It probably excludes the Three Sisters and many other landforms valued by the community, but not necessarily by professional Earth scientists. The narrow, science-centred, view of significance given by Joyce is derived from an earlier definition of a “significant geological feature’ by Legge and King (1992): “...those features of special scientific or educational value, which form the essential basis of geological education, research and reference. These features are considered by the geological community to be worthy of protection and preservation”. While a utilitarian view of significance became dominant among the official geological community, it was not the only view on offer in Australia. Sharples (1995) indicated that geodiversity elements might possess intrinsic and ecological values in addition to their utilitarian value to humans. He also noted that the heritage values (i.e. values to humans) of geodiversity included; aesthetics, inspiration, recreation, cultural development and a contribution to a ‘sense of place’ in addition to the scientific and educational values noted by Joyce (1995). This wider view of significance was adopted by the Australian Natural Heritage Charter for both biodiversity and geodiversity which gives the following definition: “Natural significance means the importance of ecosystems, biological diversity and geodiversity for their existence value, or for present and future generations in terms of their scientific, social, aesthetic and life-support value” (Cairnes 1996, p 6). Proc. Linn. Soc. N.S.w., 122. 2000 R.A.L. OSBORNE 159 Significance Criteria and Definitions It is difficult to decide how to measure or determine significance. One option is to measure significance on a scale (e.g. local, regional, national or international). Another is to define a critical cut-off level, with potential places ranking above the cut-off being significant for a particular purpose (e.g. heritage listing, reservation, consideration in planning instruments) and those below being insignificant. Significance is determined either directly by the vote of an expert panel, or by measurement against a set of criteria, usually mediated by an expert, an expert panel or a series of panels. The Register of the National Estate uses expert panels to determine significance against a set of criteria and then make a yes or no decision as to whether the place should be listed (i.e. a cut-off decision). Other systems ask experts or panels to use criteria and then rank places according to their level of significance. In federal systems of government, like Australia’s, significance assessment procedures that rank places can have serious political and financial implications. Should local government be responsible for places with local significance, state government for those with regional significance and the federal government only responsible for places with national and international significance? Since state governments run national parks, should the federal government only be responsible for internationally significant places? These questions are currently being debated in Canberra. World Heritage The International Union for the Conservation of Nature (IUCN) has the task of advising and assisting the UNESCO World Heritage Centre in implementing the World Heritage Convention. One of the main roles of the IUCN is to evaluate places nominated to the World Heritage List as having “outstanding natural value’. The process by which nominated places are evaluated is outlined by Hogan and Thorsell (2000). Article 2 of the World Heritage Convention defines natural heritage as: “natural features consisting of physical and biological formations or groups of such formations, which are of outstanding universal value from an aesthetic or scientific point of view; geological and physiographic formations...of outstanding universal value from the point of view of science or conservation; natural sites and precisely delineated natural areas of outstanding universal value from the point of view of science, conservation or natural beauty.” It is important to recognise that a standard of “outstanding universal value” is built into each part of this definition. This is a very high criterion; it is not easy to show that a natural place meets this. A key element of the process is comparing the nominated place with other similar places throughout the world. This is designed to ensure that the World Heritage List is “ only a select list of the most outstanding...from an international viewpoint”. Specific provision is made for geoheritage places in the World Heritage List. The requirement is that they should: “(a) (i) be outstanding examples representing the major stages of earth’s history,including the record of life, significant on-going geological processes in the development of landforms, or significant geomorphic or physiographic features; or (ii1) contain superlative natural phenomena or areas of exceptional natural beauty and aesthetic importance.” Proc. Linn. Soc. N.s.w., 122. 2000 160 PRESIDENTIAL ADDRESS 2000 These criteria are much more inclusive than those of the Geological Society of Australia’s concept of geological heritage, but not too dissimilar from the concept of significance given in the Australian Natural Heritage Charter. National Estate Criteria Places nominated for listing on the Register of the National Estate are evaluated against a set of eight criteria, some of which are divided into sub criteria (Australian Heritage Commission 1993). Places are ranked high, medium or low against the criteria, and then a decision is made as to whether the place does or does not meet the standard for listing. National Estate Listing is subject to both administrative and judicial review, and there are cases where both have occurred. As a consequence those involved in the evaluation process are careful to ensure that both the listing of a place and its nominated boundaries can be defended against the most rigorous inquiry. Only some of the criteria and sub criteria are relevant to geodiversity. Each of the relevant criteria, from Australian Heritage Commission (1993), are given and discussed below. Note that each criterion and sub criterion begins with the word “importance”’. It is the task of expert nominators, Evaluation Panels, Heritage Commission staff and the Commission itself to decide just how important a place must be for it to be registered. “A.1 Importance in the evolution of Australia’s flora, fauna, landscapes or climate.” Sub criterion Al is particularly applicable to geodiversity. It can include fossil localities, geological sites that give palaeoenvironmental or palaeogeographic information as well as palaeoclimate sites. Places providing evidence for plate movement could also be included. “A.2 Importance in maintaining existing processes or natural systems at the regional or national scale.” This is usually thought of as an ecological criterion, however it can just as well apply to any active Earth system of regional scale. River and groundwater systems, aeolian processes in large sandy deserts and regional longshore drift could meet this criterion. “A.3 Importance in exhibiting unusual richness or diversity of flora, fauna, landscapes or cultural features.” Geoheritage places can be rich and/or diverse. This criterion will admit both rich places with low diversity and diverse places that are not rich. Shearsby’s Wallpaper near Yass has abundant well-preserved specimens of two species of brachiopods, while the Delegate Pipes intrusions in southeastern N.S.W. contain “a large variety of rare xenolith types” (Schén 1984). “B.1 Importance for rare, endangered or uncommon flora, fauna, communities, ecosystems, natural landscapes or phenomena, or as a wilderness.” This has generally been interpreted as encompassing any natural heritage feature that is genuinely rare, endangered or uncommon. A whole range of geodiversity places have been seen to meet this criterion, including: fossil and mineral localities, outcrops of rare rock types (the olivine leucitite at El Capitan, western N.S.W.), meteorite impact lithologies (the Liddell buchite, Hunter Valley, N.S.W.), burning mountains (Mt Wingen, near Scone, N.S.W.) and unusual landforms (Australia’s only hum, a type of residual limestone hill, at Mole Creek, Tasmania). Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 161 “C.1 Importance for information contributing to wider understanding of Australian natural history, by virtue of their use as research sites, teaching sites, type localities, reference or benchmark sites.” This sub criterion has allowed a very large number of places with potential significance to be generated. There has been considerable discussion about which of these are truly significant. The reason for this is that sub criterion Cl places are an artefact of working natural scientists. A potential C1 place is created every time a scientist does field work, takes students to a specific locality, describes a new species (biological or mineralogical) with a type locality, defines a stratigraphic type section or indicates a soil reference site on a map. Most natural places that have been nominated to the Register of the National Estate have had Cl as one of a number of highly rated criteria. How to assess the significance of the large and growing number of places which rate highly simply as research, teaching, type and reference sites remains to be resolved. “D.1 Importance in demonstrating the principal characteristics of the range of landscapes, environments or ecosystems, the attributes of which identify them as being characteristic of their class.” Places that meet this criterion do not have to be rich, diverse, rare, uncommon, or used for science or teaching. They must be an outstanding example of what they are. This criterion says that the best example of something very common can be significant. It is generally seen to incorporate the concept of “representativeness”’. A representative example a feature must clearly exhibit the all, or most, of the key features of its class. This is best illustrated by a hypothetical example. Sandy beaches are very common in Australia. A representative sandy beach would have all of its components; bars, swash zone, berm and dunes intact and well developed. It would be the example of a beach you might use in a textbook. “E.1 Importance for a community for aesthetic characteristics held in high esteem or otherwise valued by the community.” This criterion solves the problem of highly regarded places that are ignored by the scientific community. The criterion talks about “a community”, which allows places valued by particular defined groups, ethnic or social also to be included. “G.1 Importance as places highly valued by the community for reasons of religious, spiritual, cultural, educational or social associations.” At first glance this might appear to be the “churches and war memorials” provision, and these places meet this criterion. Some geoheritage places have great significance to Koori people and as a consequence meet this provision. Other geoheritage places have this type of significance for Australians with a range of ethnic backgrounds. Caves in New South Wales have been used for weddings, church services and Masonic rituals (Jenolan and Wellington), dances and concerts (Abercrombie, Jenolan and Kanangra Walls), by bushrangers (Abercrombie, Cliefden, Coolah and Jenolan) and as a classroom (Wuulumin Cave). Similarly, vantage points used for ANZAC and Easter Dawn Services might qualify under this criterion: “H.1 Importance for their close associations with those individuals whose activities have been significant within the history of the nation, state or region.” A number of geoheritage places have associations with people considered significant Proc. Linn. Soc. N.s.w., 122. 2000 162 PRESIDENTIAL ADDRESS 2000 to the history of European exploration and/or the development of the natural and geological sciences both in Australia and internationally. Particular landforms are associated with or claimed to be associated with the work of early European explorers and surveyors (e.g. Thomas Mitchell and Victoria Pass and the various purported localities of Barralier’s Pass in the Blue Mountains, west of Sydney). Another strong association exists between landforms and aviation pioneers (e.g. Hargraves with Bald Hill, and Kingsford-Smith with Seven Mile Beach, both located in the Illawarra Region, south of Sydney). Examples of geoheritage places in New South Wales that have close associations with significant naturalists and Earth scientists include: PLACE ASSOCIATED PERSON/S David Moraine Edgeworth David David's Cutting, Maitland Edgeworth David El Capitan Leucitite A. Harker, Edgeworth David, Milne Curran, Etc. Fennel Bay Fossil Forest W.B. Clarke Kiama Blow Hole J.D. Dana and W.B. Clarke Mt Gibraltar, Bowral Douglas Mawson, W.R. Browne Mt Wingen (Burning Mountain) Thomas Mitchell Mt Woowoolahra Douglas Mawson Seaham Quarry Edgeworth David | Soho Street Amphibolite, Cooma W.R. Browne and Germaine Joplin Wellington Caves Richard Owen, George Cuvier, Thomas Mitchell, A.M Thomson, P. Strzelecki, G. Krefft. etc. _ Thematic Assessment and the National List One of the options currently being discussed as a replacement for the Register of the National Estate involves the development of a “National List’. The proposal is that the “National List” would include perhaps one hundred places, regarded as being significant at a national level. The ‘Re-drafted National List Criteria, version 9/11/99’, produced by Environment Australia, states that: “The National List will comprise those places, or groups of places, that are of outstanding significance for the Australia community, in that they are symbolic, exemplary or unique places reflecting the agreed themes of national importance (The National Themes). Places entered in the National List will satisfy each of the following criteria: Criterion |. the place must be a symbolic, exemplary or unique example of the highest Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 163 order in representing or demonstrating a National Theme; Judgments on the significance of a place will be tested using the sub-criteria listed below. Criterion 2. the place must have a very high level of integrity in its nationally significant values; Criterion 3. the place must possess a great capacity to demonstrate its primary National Theme, and places that in addition to this primary criterion also reflect other aspects of natural and cultural diversity will be favoured over places of equal thematic value that do not.” The sub-criteria proposed are very similar to the existing National Estate Criteria. Possible contexts for the National Themes include: “An Ancient Land’, “Continental Isolation”, “Settlement of Australia by hunting-and-gathering societies”, “European Expansion and creation of nation”, and “Encounter between cultures” (Pearson 1999). The following themes related to geodiversity are listed in the first two context areas: e “Ancient records of life and landforms. e Origin and development of biota and landforms as a result of Gondwana plate tectonics and more recent stability and long isolation. Evolution of landforms, species and ecosystems under conditions of stress. Climatic change and its impacts.” (Pearson 1999, p 18) It has been suggested that the National Themes should form the basis for promoting regional heritage tourism. The really important issues about National Themes are those concerning who develops them and on what basis are they developed. This remains to be seen. Comparison with similar places Most heritage assessment procedures require that a proposed place or item of heritage significance should be compared with similar places. In some systems this means similar places or items already listed, while in other systems it means other known similar places in the region, country or world. Fortunately, Solar System wide comparisons have yet to be considered, for if they were, basaltic volcanoes and impact craters on Earth would quickly be delisted. Comparing places, even those of the same general type, is never easy. Two of the problems that arise are: how similar do the places need to be for a comparison to be valid, and to what extent do differing regional settings add to the significance of otherwise similar places? The latter question applies to a comparison between a relict sand dune in the Blue Mountains and a dune of similar age and size in a desert region. The setting of the relict dune would make its comparison with the dune in the desert invalid. A valid comparison would be with other relict dunes, located away from modern deserts. Objections to heritage listings are often made on the basis of comparative significance. One, from a mining company, went something like: “this is not be best example of feature x, but we won’t tell you where the better examples are located”. CONSERVATION The Australian Natural Heritage Charter (Cairnes 1996) defines conservation as:”’all the processes and actions of looking after a place so as to retain its natural significance and always includes protection, maintenance and monitoring”. There are some special aspects to each of these essential components when geodiversity is being conserved. Proc. Linn. Soc. N.s.w., 122. 2000 164 PRESIDENTIAL ADDRESS 2000 Protection Protection involves using legal or social measures to ensure that the values of the place remain intact. There are a number of ways in which this can be attempted. I use the word ‘attempted’ advisedly, because legal and social measures can never be guaranteed to work. Sometimes protective measures can actually encourage destruction of, or damage to, the place they were intended to protect. Protection by secrecy There is a long history of using secrecy as a means of protecting limestone caves, fossil sites and mineral/gem localities. In the case of limestone caves this practice goes back to the early 20th century when, as legend has it, the chief guide at Jenolan Caves, Vos Wiburd, hid cave entrances by landscaping and burnt his notebooks following a dispute with his employer, the Department of Mines. Practices of this type have been undertaken by caving clubs since the late 1940s, with secret maps, restricted access publications, restricted access data bases, landscaped entrances and whispered conversations continuing to be used. The danger from management by secrecy is often not the wrong people finding out, but the proper authorities never finding out, and as a consequence failing to take appropriate action. If a secret place is really secret, then professional planners and land managers will not know about it. Local government planners will not take it into account, so it may be threatened by inappropriate development. Should the self-appointed custodians wish to take legal action to protect the place from some threat they will face the accusation that as the place is not recorded it either is not significant, or has been “discovered” simply as an excuse to stop the development. The motives of the secret-keepers may also be questioned. Those wishing to protect the caves at Mt Etna in Queensland were accused of wishing to use the caves (illegally) for their own exclusive recreation. Similar accusations could be levelled at mineral and fossil collectors with secret localities on other people’s land. This dilemma occurs when producing publications from heritage reports (e.g. Percival 1985). If the place is an open secret and it does not appear in a published list, it could be taken to indicate that it is really not so special after all. Secrets can be revealed in unexpected ways. The online version of the Register of the National Estate gives locality details for a fossil locality, followed by a condition report saying that the main threat to the place’s integrity comes from its location being more widely known. Protection by reservation It is a tradition in Australia that very important places are best protected by being placed in public ownership in a reserve or National Park. There is a long history in New South Wales of geoheritage places receiving such protection and recognition. Some significant examples are given in the following table: Rea ategea Reseda IRESERVE CATEGORY Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 165 Despite their innovative timing and promise, in most cases these reservations failed to protect the significance of the places over which they were declared. Reserves declared to protect caves, even those specifically dedicated for the “preservation of caves”, did not exclude mining (Middleton 1969) and in some cases acted to encourage it. Reserves over fossil localities usually had no trustees appointed and no bylaws to make removal of fossils illegal. In the case of the Fennel Bay Fossil Forest, reservation was a total failure. Practically all of the fossil tree stumps (estimated at 500 by Clarke 1885) have been removed, with only 30-40 remaining in 1979 (Percival 1979). National Parks, Nature Reserves and Karst Conservation Reserves offer the highest level of protection to natural heritage in New South Wales and prohibit mining. That does not mean that they offer a high level of protection to geoheritage places. The National Parks and Wildlife Act has a strong fauna and flora focus. There is no guarantee of specific management for geoheritage places and penalties for offences against non-living elements are weak. In New South Wales some geoheritage sites, particularly fossil and mineral localities, have a better history of protection on freehold land under the care of resident owners (with fences, dogs, suspicion of strangers etc.) than on public land. Protection by legal intervention Legal intervention is very expensive in both time and money and highly unpredictable as a means of protection. Legal action can usually only be triggered by an active or “real and present” threat to the place. Win or loose, the process creates polarisation and ill will, which is difficult to overcome. The legal system is often more concerned with correct process rather than environmental outcomes. Court decisions are good at stopping particular events or letting them occur, but they do not always form the basis for ongoing protection and management. As a consequence of legal action, mining ceased at Yessabah Caves near Kempsey, north coast of N.S.W., in 1991 (Osborne 1994), but the site has not been rehabilitated and the lantana continues to flourish. Protection by planning instruments ~ Local government planning instruments, such as Local Environment Plans and Development Control Plans, can be powerful tools for protecting geoheritage places of all types on both public and freehold land. Large-scale sites such as landforms and geological structures are often best protected by zoning that prevents land uses such as rural residential subdivision, which may obscure views. Small places may be protected by restrictive zonings, such as ““7J Scientific”, but this requires careful negotiation with landowners. In the current climate of corporatisation, privatisation and sale of surplus land, zoning may be the only mechanism to keep public sector landowners in check, unless the Minister decides to override local planning approval. Protection by agreement The future for a geoheritage place is often most effectively assured when its owners have entered into a conservation agreement with a State or Local Government body. This is particularly the case with small places located on rural properties. Conservation agreements can provide funding for fencing and conservation works and in some cases reductions in Local Government Rates, in exchange for an agreement to protect the place. The landowners retain their rights to control access. Resident owners frequently provide policing and management at a level not available on public sector lands. Some landowners develop long-term, sometimes multi-generational, relationships with scientists and other user groups. An example of this situation is at Cliefden Caves where two generations of landowners have maintained excellent relationships with palaeontologists and the Orange Speleological Society. Maintenance Preservation without maintenance can lead to destruction. Cairnes (1996) defined Proc. Linn. Soc. N.s.w., 122. 2000 166 PRESIDENTIAL ADDRESS 2000 maintenance as “continuous protective care”. Lack of “continuous protective care’, rather than defective legal protection, allowed the Fennel Bay Fossil Forest to be largely removed. As aconservation strategy, maintenance includes enforcement, fencing, weed control, erosion control and drainage. Enforcement does not necessarily mean patrols by rangers and security officers. It is the chance of being seen or caught that is by far the best deterrent to vandalism. The major advantage of resident landowners is being there, caring for the place, fixing the fences and applying the “heel of the owner” to the weeds. Monitoring Managers of public places set aside for conservation are required to produce Plans of Management or Conservation Plans that outline how the significant features of the place will be conserved and maintained. These plans can be comprehensive multi-volume reports or simple recipe book style documents produced to keep various levels of bosses and the interested public in their place. Sometimes few of the actions outlined in the plan take place, and without monitoring we are none the wiser. If the plan has got it wrong, the values may be destroyed, rather than conserved, in the time between the development of one plan and its successor. Even though large sums of money may be spent on producing a plan, it may not be implemented simply because the management authority has lost their copy. Monitoring does not have to be elaborate (with instruments, sensors, data loggers etc.), a simple look-see will often tell you if all is well or not. Conservation vs Use or Collection One solution to damage by humans is to limit or prohibit access or particular activities. Fencing off public areas, blocking tracks, restricting walkers to paths and gating caves are not universally popular among the outdoor recreation community. Similarly prohibiting or controlling collecting will quickly raise the ire of lapidaries, fossil collectors and some professional educators and scientists. Four wheel drive enthusiasts, trail riders (bike and horse), bush walkers, teachers, youth leaders, rock climbers, cavers, ecotourism operators, respectable members of this Society and many others all want to be able to do their thing, because it is always someone else who does the damage. As a consequence land managers often do their duty at some peril. Ownership by discovery One of the most common and fallacious arguments facing land mangers arises from the assumption that those who discover something own it, are entitled to unrestricted use of it, or should determine how it is used. The notion of ownership by discovery is found among palaeontologists, fossil and mineral collectors, and is particularly prevalent among cavers. Discoverers often view those with legal ownership and/or responsibility for management of their discovery with distain and suspicion. “I found it, what right do they have to tell me what to do”, 1s a view frequently expressed. Should anyone use/access? If something is significant and really fragile, perhaps people should be kept away completely, no matter who they are or what they wish to do. This approach can vary from forcing people to view the feature from afar to entirely preventing access. People may question the value of something they are not able to directly experience. Alternative approaches have been developed which allow a visitor experience while keeping people away from the feature itself including building an artificial replica adjacent to the real feature (as at Lascaux Cave, France), exhibiting photographs and models of the feature and using film, video or computer technology to produce a virtual experience. Who should use/access? [f itis decided that some people will be allowed in and most will be excluded, there needs to be a proper rationale for doing so. Allowing some people access or use on the Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 167 basis of merit is a guaranteed way to promote disputes. Bona-fide researchers are often given privileged access, but can this always be justified as beneficial to conservation and management? Where significant features occur in areas used for commercial tourism there is a simple (but not necessarily socially equitable) solution. Make access to the most fragile features expensive and thus keep visitation down. This occurs at Jenolan Caves. If a vulnerable place is a public asset, such as a National Park or reserve, how can the access or use privileges of a particular group be favoured against those of the population at large? Access and use privileges in Australia have frequently centred on questions of merit. Real scientists, walkers and members of accredited rock climbing and caving groups have been the winners. Amateurs, people in cars, parents with children in strollers and competent adventurers who don’t join clubs (or belong to the wrong ones) have often been excluded. From a conservation and management perspective the only relevant questions are not who the prospective users are, but what their impact will be on the place and will they be able to undertake the activity without unacceptable risk to their own or public safety. This is not always a popular view. In the U.S. National Parks a ballot system is used to determine who is able to undertake some over-popular treks, and in Western Australia access to some delicate caves 1s limited to a fixed number of visits in the applicant’s life. These systems solve some of the problems inherent in controls based on merit. Should anyone collect? If the Fennel Bay Fossil Forest was found today we probably would not allow the petrified logs to be used as railway ballast or fencing materials. A land manager today would take their responsibility to keep the site intact seriously. Studies of collected specimens may greatly enhance understanding of the place, with benefits to management and interpretation. On the other hand, advances in technology may make some forms of collecting obsolete in the near future. High quality imaging, 3D rendering and lightweight portable instruments for chemical and mineral analysis are already reducing the importance of the hand specimen. A thoughtful manager might say to a researcher; “come back when you no longer need to collect”. Following well-known disasters like the extinction of the Dodo, biologists have developed ethical collecting protocols. At a basic level these are that you don’t collect the only living specimen and you don’t collect so much of a population as to threaten it’s survival. Earth scientists rarely give consideration to ethical collecting. In my field of research, working in heritage-listed caves, the issue of ethical collecting is never far away. Micro sampling, indirect sampling and ensuring that excavations leave stratigraphic sections intact, are the orders of the day. There have been geological collecting events that have verged on the dodesque! In the 1930s the Australian Museum collected over one thousand specimens of stalactites, stalagmites, helictites and crystal clusters from Cliefden Caves in order to construct an exhibit (Hodge-Smith 1936). Issues to be considered in managing collecting include: ensuring a sufficient range and quantity of material is left intact for future research managing and limiting collateral damage from collecting ensuring that the amount collected is not greater than is really necessary ensuring that non-collecting methods are considered, before collection takes place deciding whether the best specimen should stay in situ, or be moved to a museum. Who should collect, how and what should they take? Where the significant features are abundant and their survival in situ is unlikely there is no need to control collecting. Mulbring Quarry in the Hunter Valley of N.S.W. exposes highly fossiliferous siltstone, used for road metal. In the normal course of events Proc. Linn. Soc. N.s.w., 122. 2000 168 PRESIDENTIAL ADDRESS 2000 the fossils will be broken and compacted into roads. Continuing quarrying operations expose more fossils, acting as a form of self-management (Stevenson 1981). While there is no conservation reason to control access or collecting, public safety, security and liability issues need to be considered. When a resource is scarce, collecting may be the greatest threat to its survival. Placing reference specimens in collections, however, may be the geoheritage equivalent to keeping threatened species in zoos. It is reasonable to argue that only specialists should undertake this type of collecting and that very good reasons need to be given to justify additional collecting. Land managers who treat requests from intending scientific collectors with suspicion do so with the benefit of hindsight. The history of vertebrate fossil collecting from New South Wales caves (Osborne 1991) includes examples where leading researchers removed deposits in their entirety, made no stratigraphic observations and kept no proper records of provenance. Much of the “cart loads” of bone in museums collected during the 19th century are of little value. Some collection sites can’t be reinvestigated because there is nothing left. Modern controls on collecting must ensure that collection of fossils for taxonomic studies, for instance, does not make future stratigraphic or palaeoecological studies impossible. Collection based studies must be able to justify the damage done to the site by collection on the basis of tangible benefits to management and interpretation. Where should the collections go? Collecting does not cease to be an issue when the rock, fossil or mineral is removed from the ground and carefully packaged for transport; in fact some of the most complex and intractable issues are just beginning. The first issue, which must be resolved, but often isn’t, is who owns the specimen? Collectors, both amateur and professional, frequently assume that once they dig it up and write an institutional specimen number on it, that they, or their institution, are the owners. This is usually not the case. Most often the specimen remains the property of the landowner or managing state or local government authority; the exception is where statutory collecting rights exist, eg Geological Survey staff. Whoever owns the specimen has the right to decide what should become of it. There are a number of issues to be considered in making such a decision: can the specimen be wholly or partly destroyed, or must it be kept intact? should the specimen be preserved or disposed of at the end of the current study? if the specimen is to be disposed of, can it be destroyed, sold, swapped or gifted? if the specimen is to be preserved where, by whom and under what conditions? When it is decided that the specimens should be preserved, the issues of where and by who can become complex and emotive. There are a number of worthy, competing alternatives that need to be considered: significant specimens should be housed in state or national institutions specimens can be housed in overseas institutions and at a range of teaching and research institutions, giving status and recognition to the place all specimens should be housed in a repository at the site type specimens should be housed in state or national institutions, all others should be returned to the site and housed in a repository at the site ° the specimens should become the property of the appropriate state collecting institution ° all specimens should remain the property of the owner/management authority of the place, specimens not on site will be considered to be on loan. There are good arguments for and against all these propositions. Whatever is decided, much angst will be avoided if clear decisions are made at the outset. Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 169 ACTIVE PHYSICAL INTERVENTION Many places are best conserved by doing very little, but in some cases there is a need for quite substantial intervention. Regeneration Regeneration involves allowing natural processes to restore something of significance. It is most appropriate for conserving partly disturbed living systems where regrowth and reproduction can, over time, repair the damage. Partly disturbed active landforms such as beaches and dunes do have a capacity to regenerate, as do some (but not most) constructive chemical deposits (rim pools, tufas and some speleothems). The significant features of most geoheritage places, however, don’t regrow or reproduce. Stabilisation (Preservation) This involves enhancing the physical strength or resistance of the significant features to weathering and erosion. It is a form of preservation because it attempts to slow or stop natural processes. Stabilisation can involve a range of techniques such as sealing, impregnation, grouting, rock bolting, reinforcing with rods, physically supporting etc. The main problem with these types of interventions 1s that once installed they require perpetual maintenance. Without maintenance the intervention may often cause more long-term damage than would have otherwise occurred. Use of chemically unstable sealants or steel rods that rust, can result in problems that require expensive remediation. Hardening (Preservation) Hardening is one of the most effective ways to preserve places from damage such as wear, trampling and breakage, caused by high levels of visitation. Typical hardening measures include concrete paths, rails and protective fencing, which increase resistance to the effects of people. Poorly designed or installed hardening can be intrusive and detrimental to the visitor experience. Hardening is the only option if visitation exceeds a few thousand per year or the place is easily damaged. Hardening has been standard practice at fragile places such as show caves, but is becoming more common at places that attract large visitor numbers. Substantial hardening has been undertaken at North Head, Sydney Harbour, to prevent erosion and trampling. Scaling (Restoration) Bedrock features exposed in artificial outcrops are frequently obscured by weathering and slope debris. Cleaning or scraping back the surface of the outcrop can reveal the significant features. Scaling, as a restorative activity, should be distinguished from scaling for public safety/geotechnical purposes, which, while an essential management activity, may threaten the significance of the place. Re-exposure (Restoration or Enhancement) Re-exposure involves removing more than a small amount of obscuring dust or debris from a feature. It can be restoration if the obscuring mantle is a result of a recent rockfall, or enhancement if the obscuring material has been in place for a considerable time. While re-exposure may enhance the view of a feature, it may make it more vulnerable to weathering and erosion and other natural elements may be degraded in the process. Proposals to re-expose a site must be carefully evaluated and not undertaken lightly. Short-term advantages of improved views need to be weighed against increased maintenance and possible reductions in life expectancy of the feature. Proc. Linn. Soc. N.s.w., 122. 2000 170 PRESIDENTIAL ADDRESS 2000 Re-burial (Preservation) Some features are preserved best by being buried, or re-buried. This is the case with features exposed through excavation. For re-burial to be considered, a feature must be so significant that its preservation outweighs the need for it to be seen. Features must also be more likely to survive under an artificial mantle of earth than at the surface. Re-burial is rarely used and does not always have the desired effect. The hominid footprints at Laetoli, Tanzania were re-buried, but were later exhumed and re-buried again following damage by the roots of trees growing in the earth covering the site. Protective salvage (Preservation) Protective salvage is removing significant material from a place to protect it from destruction or damage from imminent natural or human causes. Protective salvage is most often used to remove fossils from danger. Alex Richie (Australian Museum, Sydney) has been involved in a number of salvage operations including recovering fish fossils at Eden, south coast N.S.W., before they were destroyed by natural retreat of a sea cliff and at Somersby, near Gosford, N.S.W., where they were exposed in an active quarry. Protective salvage agreements, such as that at Somersby, can be made with quarry operators, but are difficult to arrange. There must be a high level of trust between the operator and those involved in salvage and trained personnel must available on call carry out the work quickly. Unfortunately much 1s lost because operators feel it is too dangerous to the continuation of their operation to report interesting material that they may unearth. Reinstatement Reinstatement is putting something back into the environment that was once there, but is now missing. Most bush-regeneration projects are actually reinstatement. Proposals to clone mammoths and thylacines are extreme examples of reinstatement. Reinstatement is rarely, if ever, appropriate in geoheritage places. Initially, the only example I could think of was replacing broken stalactites using araldite and splints, but better and larger scale examples are the artificial sand dunes constructed behind surf beaches along the New South Wales coast. INTERPRETATION Interpretation involves building a bridge between a place and those that visit and manage it. We are apt to think that visitors are the main audience for interpretation, but unless owners and managers understand and value places in their care, the chances for long-term conservation are poor. Lack of community knowledge The main problem confronting geodiversity interpretation is a lack of community knowledge and understanding. While many in the community have some understanding of elementary ideas in biology, ecology and biodiversity conservation, there is very little community understanding of the basic ideas of Earth science. One reason for this lack of information is a lack of accessible literature. It is relatively easy to obtain popular information about local flora and fauna. There are many general books and a number of specific guides, particularly to regional flora. There are very few comparable publications about rocks, landforms and soils. Similarly, interpretation material produced for National Parks, and programs run by environment centres, visitor centres and field study centres, almost exclusively focus on the biological, and are usually produced and managed by staff without much knowledge of geology. This does not mean that there is a lack of public interest in geodiversity, just that there are few mechanisms for engaging that interest. It is difficult to convince editors, producers and teachers that Earth features and processes (with the possible exception of Proc. Linn. Soc. N.s.w., 122. 2000 R.A.L. OSBORNE 171 dinosaurs, earthquakes and volcanoes) are interesting, or worth the risk, when sharks, killer whales and cuddly animals have well-established, and regenerating markets. Research for Interpretation The lack of mainstream scientific interest in geoheritage places with general public interest has created the need for applied basic research to provide a basis for interpretation and to answer questions frequently asked by the public. Questions raised by interpretation are often complex and multidisciplinary. They do not lie within conventional disciplinary research programs, nor are they likely to be answered by industry-based applied research. Much of my research at Jenolan, Wellington and Wombeyan Caves has been directed towards answering two questions frequently asked by visitors; “how old are the caves?” and “how did they form?” There have been numerous attempts to improve geodiversity interpretation and education in New South Wales (Osborne 1992; Wilkins and Osborne 1996). Making worthwhile and lasting progress in this area remains one of the greatest challenges for the future. CONCLUSIONS Practicing geodiversity requires a range of skills and an approach to the Earth sciences not frequently found among academic or professional geoscientists. Expansion of work in geodiversity will largely depend on changing the attitude and focus of politicians and nature/heritage conservation policy-makers. The introduction and adoption by some Local Government organisations of the Australian Natural Heritage Charter is a significant move in this direction. Geodiversity has the potential to provide a whole new sphere of employment for Earth science graduates. For this to occur there will need to be a change not only in the content of their training, but also in the values and attitudes instilled in them. The time for regarding Earth scientists working in heritage conservation as traitors to the profession has long since passed. ACKNOWLEDGMENTS It’s a little over twenty years since I first became involved in geoheritage conservation. This has not made me many friends among the academic and professional geological community, and one senior academic considered my work as “not in the national interest”. I am therefore most appreciative of all those who have been supportive, in particular: My former colleagues on the New South Wales Geological Sites and Monuments Subcommittee of the Geological Society of Australia: Betty Collett, Pat Conaghan, Hugh Henry, Bob Jones, Ian Percival, the late Richard Sch6n, and Michael Williams, who worked with me for almost 13 years to identify, document and conserve a wide range of geoheritage places. Paul Adam, Lorraine Cairnes, Roger Carolyn, Martin Denny, George Gibbons and Dominic Sivertsen who served with me on the New South Wales Natural Environment Evaluation Panel of the Australian Heritage Commission, supported geoheritage issues and helped incorporate them into the mainstream of heritage conservation. Sydney Craythorne, Elery Hamilton-Smith, Patrick Larkin, Grant Gartrell, Ernst Holland, Andy Spate, Kier Vaughn-Taylor, and Dianne Vavryn, karst specialists, managers and conservationists who continue to make major contributions to geoheritage conservation. Lyn Sutherland, Ross Pogson, Alan Jones, Alex Ritchie, and Gail Webb, colleagues at the Australian Museum for their continuing assistance and support and Des Griffin, former Director, Australian Museum, for his unswerving support in difficult situations. Meg Switzer, Roland Eberhard, Nathan Wales and Cameron Slatyer of the Australian Heritage Commission and Environment Australia. Pavel Bosak and Vaslav Cilek of the Geological Institute, Prague and Daniel Rozek of the Institute for the Conservation of Natural and Cultural Heritage, Nova Gorica, overseas colleagues in geoheritage, who provided a different perspective on the subject and continue to share their time and resources generously. Proc. Linn. Soc. N.s.w., 122. 2000 172 PRESIDENTIAL ADDRESS 2000 Manly Council, Pittwater Council, Shoalhaven City Council and Wellington Council, which have taken geodiversity seriously and supported identification and documentation projects. Penney, my wife and partner in geodiversity, has not only been supportive, but has participated in fieldwork, trudged through wetlands (swamps) in the dark, and commented on turgid management plans, consultants reports, heritage nominations and this paper. A NOTE OF THE LITERATURE The references include a number of works from the “grey” literature as well as conventional books and journals. Much of this material is held in the libraries of the NSW Department of Planning, NSW National Parks and Wildlife Service, NSW Department of Mineral Resources, and Environment Australia (Canberra). Copies of unpublished consulting reports are generally available from the commissioning agencies. NOTE ADDED IN PREP There have been a number of significant developments in the political environment of geodiversity conservation since the Presidential Address in March 2000. In April, the Office of the Sydney Harbour Manager launched the “Spectacle Island Statement for Conserving the Natural Heritage of the Sydney Harbour Catchment”. This six-page document contains a statement on the geodiversity of Sydney Harbour. In July, the final meeting of the NSW Natural Environment Evaluation Panel of the Australian Heritage Commission was held. This probably marks the beginning of the end of both the Register of the National Estate and the Australian Heritage Commission. Public briefings were held in August to explain the proposed new Commonwealth approach to heritage and the National List. Legislation is apparently to be placed before Federal Parliament in 2001. The future of geoheritage identification and documentation in New South Wales looks bleak unless the NSW Heritage Council and/or the National Parks and Wildlife Service (or some other body) takes up the role formerly played by the Australian Heritage Commission. REFERENCES Anon, (1948). 48/424. Lime deposits near Dungog, Gloucester and Grafton in Limestone Deposits of the North Coast. New South Wales Geological Survey Report GS 1952/031. Australian Heritage Commission, (1993). ‘Diversity Assessment and the Identification of National Estate Places’. Regional Assessment Branch, Australian Heritage Commission, Canberra. 13 p. Cairnes, L.B., (1996). ‘Australian Natural Heritage Charter: Standards and Principles for the Conservation of Places of Natural Heritage’. Australian Heritage Commission, Canberra, 18 p. Carne, J.E. and Jones, L.J., (1919). The Limestone Deposits of New South Wales. Geological Survey of New South Wales, Mineral Resources 25, 411 p. Clarke, W.B., (1885). Awaba fossil forest. Annual Report of the New South Wales Department of Mines 1884, 156-159. David, T.W.E., Taylor, T.G., Woolnough, W.G. and Foxall, H.G., (1905). Occurrence of the pseudomorph glendonite in New South Wales. Records of the Geological Survey of New South Wales 8, 161-179. Dixon, G., (1996). Geoconservation: An International Review and Strategy for Tasmania. Parks and Wildlife Service Tasmania, Occasional Paper 35, 1-101. Eberhard, R., (1997). (Ed.) ‘Pattern and Process: Towards a regional approach for National Estate Assessment of Geodiversity’. Environment Australia, 1997 Technical Series No 2. Canberra, 102 p. Proc. Linn. Soc. n.s.w., 122. 2000 R.A.L. OSBORNE 173 Hogan, R. and Thorsell, J., (2000). “Guidelines for the External Review of Natural World Heritage Nominations’. International Union for the Conservation of Nature, Gland (Switzerland). 13 p. Hodge-Smith, T., (1936). A limestone cave in the Museum. Australian Museum Magazine 6(2), 39-46. Jaquet, J.B., (1901). The iron ore deposits of New South Wales with maps, plates and sections. Memoirs of the Geological Survey of New South Wales, Geology 2, 66-69. Jaquet, J.B. and Harper L.F., (1899). “Geological map with section of the country in the vicinity of the Williams and Karuah rivers north of Port Stephens, showing associated beds of magnetic ironstone’. Department of Mines and Agriculture, Sydney. [Distributed in back pocket of Memoirs of the Geological Survey of New South Wales, Geology 2] Joyce, E.B., (1995). ‘Assessing the Significance of Geological Heritage: A methodology study for the Australian Heritage Commission.’ A report prepared for the Australian Heritage Commission by the Standing Committee for Geological Heritage of the Geological Society of Australia Inc, 19 p + appendices. Legge, P. and King, R.L., (1992). Geological Society of Australia Inc. Policy on Geological Heritage in Australia. The Australian Geologist 85, 18-19. Lishmund, S.R., Dawood, A.D. and Langley, W.V., (1986). The Limestone Deposits of New South Wales. Geological Survey of New South Wales Mineral Resources 25 2nd Edition, 373 p. Middleton, G.J., (1969). The case for the conservation of Colong Caves Reserve, New South Wales, Australia. Studies in Speleology 2(1), 1-11. Osborne, R.A.L., (1991). Red Earth and Bones: The History of Cave Sediment Studies in New South Wales, Australia. Journal of Earth Sciences History 10(1), 13-28. Osborne, R.A.L., (1992) The earth sciences, schools and beyond. ‘Sydney Universities Consortium of Geology and Geophysics Symposium on Geology and the Community *, 17-18. Osborne, R.A.L., (1994). Caves, cement, bats and tourists: karst science and limestone resource management in Australia. Journal and Proceedings of the Royal Society of New South Wales 127, 1-22. Osborne, R.A.L., (1996). ‘Tapalla Point Geological Site, Huskisson’. Report to Shoalhaven City Council, January 1996. 10 p. Osborne, R.A.L., (1997). The regional context for assessing heritage values of geodiversity. In: Eberhard, R. (Ed.) “Pattern and Process: Towards a regional approach for National Estate Assessment of Geodiversity’. Environment Australia, 1997 Technical Series No 2. Canberra, 9-14. Osborne, R.A.L., (1998). “Karst of the New England, Stage 1, NEGP Project NEP 95 321: Karst of the eastern New England, New South Wales’. Report prepared for the Australian Heritage Commission and the Department of Urban Affairs and Planning, New South Wales, 3 volumes. Osborne, R.A.L., Docker, B. and Salem, L., (1998). ‘Places of Geoheritage Significance in New South Wales Comprehensive Regional Assessment (CRA) Forest Regions.’ Report to Environment Australia. 41 p + spreadsheets. Osborne, R.A.L. and Osborne, P.J., (2000). ‘Geoheritage significance of the Elizabeth Street Faults, Intersection of Cabarita Road, Wanawong Road, Elizabeth Street and Patrick Street, Avalon.’ Report to Pittwater Council, 8 p. Pearson, M., (1999). “A National List System”. Discussion Paper for the Australian Heritage Commission, 19th August 1999, 31 p. Percival, I.G., (1979). “The Geological Heritage of New South Wales’. Report prepared for the Australian Heritage Commission and the Planning and Environment Commission of New South Wales, 277 p. Percival, I.G., (1985). ‘The Geological Heritage of New South Wales, Volume 1’. National Parks and Wildlife Service, Sydney, 136 p. Roberts, J., Engel, B., Lennox, M. and Chapman, J., (1991). ‘Dungog 1:100,000 Geological Sheet 9233’. New South Wales Geological Survey, Sydney. Schon, R.W., (1984), “The Geological Heritage of New South Wales, Volume 3’. Report prepared for the Australian Heritage Commission and the New South Wales Department of Environment and Planning, 216 p. : Semeniuk, V., (1997). The linkage between biodiversity and geodiversity. In: Eberhard, R. (Ed.) ‘Pattern and Process: Towards a regional approach for National Estate Assessment of Geodiversity’. Environment Australia, 1997 Technical Series No 2. Canberra, 51-58. Sharples, C., (1993). “ A Methodology for the Identification of Significant Landforms and Geological Sites for Geoconservation Purposes’. Report to Forestry Commission, Tasmania. Sharples, C., (1995). Geoconservation in forest management-principles and procedures. Tasforests. 7, 37-49. Stevenson, B., (1981). ‘The Geological Heritage of New South Wales Volume 2’. Report prepared for the Australian Heritage Commission and the New South Wales Department of Environment and Planning, 202 p. Wilkins, C. and Osborne, A, (1996). Interpretation of geological heritage in New South Wales. In: Facer, R.A. (Ed.). “Geology and the Community; Ninth Edgeworth David Day Symposium’. Earth Resources Foundation, University of Sydney, 51-58. Willis, P.M.A., (1993). ‘Vertebrate (Tetrapod) Palaeontological Sites in New South Wales’. National Parks Association of N.S.W. Inc., 225 p. Proc. LINN. Soc. 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Ai / eal pial adds ate Pv etmaw: Lig Gal sons Lavoro hice Motegi ciate, 1 ewe: pay) aie? r hy nh U cries Way y anhwrts. er es ERRATA Some copies of Volume 121 contained figures that were not well defined. The following figures can be substituted for figures from Volume 121 as indicated. They have been reprinted to the same size as they appeared in volume 121 and can therefore be inserted directly over them. The following three pages are from: McAlpine, D.K. (1999). Australian signal flies of the genus Rhytidortalis (Diptera: Platystomatidae). Replacement Figures | and 2, page 149 Figures 1, 2. Antennae of Rhytidortalis averni: 1, male; 2, female. Proc. Linn. Soc. N.s.w., 122. 2000 | ee aa as é gnivolich iT. banish iow ton sis age wusab banisingo. 1st cme eokgon: head sybd yd osecibni 2a (C1 simyih mont 2omgit 108 hetaaedae 3d £ hensani od stotetsd! aso bas 4 Cf samioy of bemsdgs: yaillae asin oone ont of D BO 9 a en Siti Rida: ott oa ei. ~. : j (aoe = ie: See otemolS ohana { are ssc 19 ‘oats ty Se ues = rn { haa t woe Sek vw mud page 150 ’ Replacement McAlpine (1999) Figures 3 & 4 Figures 3, 4. Left mesopleural region of Rhytidortalis averni: 3, male; 4, female. Proc. Linn. Soc. N.s.w., 122. 2000 3 £ “ae < z 3° te B44 8 i & e A es stosra) + Lalor. Wreormn silent Jo noigey leantepeaonr fis) ff vonrgit Replacement McAlpine (1999) Figures 12-16, page 162 \ Figures 12-16. Rhytidortalis averni: 12, surface of mesoscutum showing zones devoid of pruinescence (visually black spots) x 176; 13, dorsal view of female tergites 4 and 5, spiracles inducated x 55; 14, distal part of aedeagus X 108; 15, base of arista x 510;16, male genital complex from left x 58. PEN and INK change. McAlpine (1999) page 167 line 6 (under Rhytidortalis kelseyi) should read: Description © o unknown) Proc. Linn. Soc. N.s.w., 122. 2000 ERRATA Some copies of Volume 121 contained figures that were not well defined. The following figures can be substituted for figures from Volume 121 as indicated. They have been reprinted to the same size as they appeared in volume 121 and can therefore be inserted directly over them. Clague, C.I., Coles, R.B., Whybird, O.J., Spencer, H.J. and Flemons, P. (1999). The occurrence and distribution of the tube-nosed insectivorous bat (Murina florium) in Australia. Replacement Figure 2, page 183 (a) capture records (b) acoustic detection records Figure 2. (a) capture localities (n=11) of Murina florium for wet tropics sites listed in Table 2, shown as filled dots. Gray area is the total predicted distribution for M. florium using a BIOCLIM model based on the seven climate parameters listed in Table 5 for capture sites only. (b) acoustic detection localities (n=14) of M. florium, details as in (a). Predicted distribution applies to acoustic detection sites only. For further details see Fig.1. Proc. Linn. Soc. N.s.w., 122. 2000 | gah 0001). a, Aotiatt bas tat. wets “UO: tnigtW | _ Wi To (MGs) antiifeae! pottaetsh alison iy -Spioltaints | _ i Kise hence eae sr pail b wight > paded avec att! baieoibat Pathe gen poate aot pada’ od, ole ai oes hie — . — er oa cece anu), bik i atlanta ted eumeuaal = to - slits sn aoeue © old nal este aig Yaw 40 in Aa ian af ovo boned ybomt MLISOWA s nine |. grt soe elennade vant 9977 io Sole Oot 5) ae wk mela es if a ae : 7 7 p Replacement Clague et al. (1999) Figure 1, page 181 Cape Bedford Cooktown Murine florium sites Mareeba Innisfail ~ Hinchinbrook Island Palm Islands Cape Cleveland Townsville 147° Figure 1. Map of all recorded sites (n=25) for Murina florium in the wet tropics region of Australia (listed in Table 2) shown as filled dots. Grey area is the total predicted distribution for M. florium using a BIOCLIM model based on the seven climate parameters listed in Table 5 for all localities. The BIOCLIM climate model covers a land surface area (including islands) bounded approximately by the mape (Cape Bedford to Cape Cleveland) and up to 120 km inland. This area contains the wet tropics region of Australia as defined by Nix and Switzer (1991). Proc. Linn. Soc. N.s.w., 122. 2000 mi feteil) siletenA Ww hots tabrwo Mi1. PHOS 5 yeti. Sarva arcane saber re nail reli aa al open (1001) tsaitw ieee woes MONG S01, a CT = ae 7 ERRATA Some copies of Volume 121 contained figures that were not well defined. The following figures can be substituted for figures from Volume 121 as indicated. They have been re- printed to the same size as they appeared in volume 121 and can therefore be inserted directly over them. Meek, P.D. and Triggs, B. (1999). A record of Hastings River mouse (Pseudomys oralis) in a fox (Vulpes vulpes) scat from New South Wales. Replacement Figure 1, page 194 Marengo State Forest ees een Figure 1. New South Wales showing the location of Marengo State Forest. Proc. Linn. Soc. N.s.w., 122. 2000 / Bahweatl 2d 3 j owewd ues bos 181 —. er | ee oT F bonita tow i fon ow Pee oe ene {§ - saat Sr Soudeed - -o1 nogd saved: betentbat an ff1 senmnhoy sos} et ae ae (an pone) patio a9vift Bais Ww. pai ke (0001), a Per - Racal thuo® weV' son Hi00 6 ee say Aetaa, 1 sat cas i - OO0f [fi wk cot ent soni a ERRATA An error occurred in Table | of the paper by Smithers et al. on page 111 in Volume 121. The next page contains a replacement for that figure. Smithers, C.N., Peters, J.V. and Thornton, I.W.B. (1999). The Psocoptera (Insecta) of Norfolk and Philip Islands: occurrence, status and zoogeography. Proc. Linn. Soc. nN.s.w., 122. 2000 ie ( i) Wnt £51 cay om el owt Table 1. Psocoptera recorded from Norfolk and Philip Islands. SPECIES (E = Species probably endemic to NORFOLK ISLAND PHILIP ISLAND Norfolk/Philip Islands - 11 species) Earlier In 1998 Earlier In 1998 Pea | [egvclaas iced OS Va EO ee a Ee ee ae ewumnumerantinesnn won EX Xe Xo RE EDM | Kd |Pieroxanium ralstonae EX eae MEROGUDAE See ee PERT S| Coon poe SS EEE Ea Panna Pay Trace kts ees ore Lingumeremonm =< ET | eX kX KC RECTEIUSID AE eee |e es eee | Celie ai SS a Eee Ee ae Ea BECHORS OCI NE Seinen 2 Skee oe ee el ee [Eeigince iin ie on eae en ee es i, ae eae ae REciopnocisiinoniatss mat eames | exer ee REcionnoausnichandsiv es sen (cmon se tea eee IEERIBSOCIDE Ne ee RESEUDOCAE CH MIDAE Uso eis WS a ee Pucierocaccilins vaniabilis oe nls xe | [NCIC PSOUCI USS ee a le [Ae TE el Ae IPHIROTARSIDAE RPE e en ele es at aos [kino dialin aii ae ES ae Ee ae ESOCID AE Ran ee ce [Mia iar en Ae one ae | EVNOnSOCIDAES er en a es ee [RAO MSTA ge ea rel eC | Miia aims SC SES TT a SS OSes nics lotaliiumberonspecies ume a ms ee nue i P<] P<] P< Proc. Linn. Soc. N.s.w., 122. 2000 aoe ee en eo 7 AC AO x _ Xk atone nits | eee - Saar Sar et OS a ae DA MER aan Gratoia = aaioag2 102 ___ eaivage 10. 130ra The Linnean Society of New South Wales publishes in its proceedings original papers and review articles dealing with biological and earth sciences. Intending authors should contact the Secretary (PO Box 137, Matraville NSW 2036, Australia) for instructions for the preparation of manuscripts and procedures for submission. Instructions to authors are also available on the society’s web page (http://www.acay.com.au/~linnsoc/welcome.html). Manuscripts not prepared in accordance with the society’s instructions will not be considered. PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W. VOLUME 122 Issued 22 December 2000 CONTENTS 1 AMANDA REID Eight New Planipapillus (Onychophora: Peripatopsidae) from Southeastern Aveta 33 —_‘(I.D. LINDLEY Pentremites australis sp. nov., a New Lower Carboniferous (Tournaisian) Blastoid from New South Wales 43. W.B. KeiTH HoLmes The Middle Triassic Megafossil Flora of the Basin Creek Formation, Nymboida Coal Measures, NSW, Australia. Part 1: Bryophyta, Sohenophyta. 69 M.A. Hancock, B.V. Timms, J.K. Morton AND B.A. RENSHAW The Structure of the Littoral Invertebrate Communities of the Kosciuszko Region Lakes 79 _TSUYOSHI KOBAYASHI, SIMON WILLIAMS AND AMANDA KOTLASH Autotrophic Picoplankton in a Regulated Coastal River in New South Wales 89 Lyn A. BEARD AND GORDON C. GRIGG Reproduction in the Short-beaked Echidna, Tachyglossus aculeatus: Field Observations at an Elevated Site in South-east Queensland. 101 ~Leanne Armano, W. D. L. Ripe AND GRAHAM TAYLOR The Stratigraphy and Palaeontology of Teapot Creek, MacLaughlin River, NSW 123 U. Karsten Occurrence of photoprotective mycosporine-like amino acid compounds (MAAs) in marine red macroalgae from temperate Australian waters. 131. L.T. AbLem anp B.V. Timms Peay ereeiy of the freshwater Peracarida (Crustacea) from Ben Tops, N 143 J.W. Douc.as anp P. BRown | Notes on Successful Spawning and Recruitment of a Stocked Population of the Endangered Australian Freshwater Fish, Trout Cod, Maccullochella macquariensis (Cuvier) (Percichthyidae) 149 R.A.L. OsBorRNE Presidential address 1999/2000 Geodiversity: “green” geology in action ERRATA from Volume 121 Printed by Southwood Press Pty Ltd, 80-92 Chapel Street, Marrickville 2204 wih ni 5 WHSE 05848 Mere ee ee vet 2% Sooke eet SS Beene wien te Ss meen - Sines ee eh aninasl MYR Geeta gah win War Samaras quien sey nae spiced r steak ON Sue A Anwernine acura! ean tte gt “\ * SUN FA te Eareerres eye a Sab tn earner ety, weve ay gy ieee VEN wsietcad, Jeremy sete pony Ledeen hate eenessateed we WOE NERY ela oe Garannce Cerne 8 AE ERD Ra as gin Ser A WNsa ee wey Serr ees PATRON ee SAO RENARD a oes wg SLAPS DAS “Sages, Novay MPABM pws nay MO ARNEC Eh Oe ete Oh SURE SAVES AN NAVARRA st TTT SE MEAS Seow APRON aan ep a # ake ey SES. 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