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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.
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sperm transfer 24, 1517-1527.
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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
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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
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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
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Proc. Linn. Soc. N.s.w., 122. 2000
84 AUTOTROPHIC PICOPLANKTON IN A COASTAL RIVER
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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
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Paerl, H.W. (1977). Ultraphytoplankton biomass and production in some New Zeland lakes. New Zealand
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Weisse, T. (1988). Dynamics of autotrophic picoplankton in Lake Constance. Journal of Plankton Research
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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.
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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
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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.
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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
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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
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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
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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
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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.
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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
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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
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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).
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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.”
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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).
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“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
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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
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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.
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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
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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
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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
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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
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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.
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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.
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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
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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
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Cairnes, L.B., (1996). ‘Australian Natural Heritage Charter: Standards and Principles for the Conservation of
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Carne, J.E. and Jones, L.J., (1919). The Limestone Deposits of New South Wales. Geological Survey of New
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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. N.s.w., 122. 2000
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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
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Figures 3, 4. Left mesopleural region of Rhytidortalis averni: 3, male; 4, female.
Proc. Linn. Soc. N.s.w., 122. 2000
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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
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Cape Bedford
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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
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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
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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
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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
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MEROGUDAE See ee PERT S|
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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
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IEERIBSOCIDE Ne ee
RESEUDOCAE CH MIDAE Uso eis WS a ee
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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
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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
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