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VOLUME 131 July 2010
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The Middle Triassic Megafossil Flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales, Australia. Part 8. The Genera Nilssonia, Taeniopteris, Linguifolium, Gontriglossa and Scoresbya
W.B. KeitH Hoitmes', H.M. ANDERSON? AND J.A. WEBB?
'46 Kurrajong Street, Dorrigo, NSW, 2453, Australia (wbkholmes@hotmail.com). Hon. Research Fellow, University of New England, Armidale, NSW. ? 46 Kurrajong Street, Dorrigo, NSW, 2453 Australia. Hon. Palaeobotanist, South African Biodiversity Institute, Pretoria 0001 South Africa. 3Environmental Geoscience Department, La Trobe University, 3086, Victoria.
Holmes, W.B. K., Anderson H.M. and Webb, J.A. (2010). The Middle Triassic Megafossil Flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales, Australia. Part 8. The Genera Nilssonia, Taeniopteris, Linguifolium, Gontriglossa and Scoresbya. Proceedings of the Linnean Society of New South Wales 131, 1-26.
Ten taxa of simple leaves in the genera Nilssonia, Taeniopteris, Linguifolium and Gontriglossa and a lobed leaf in the genus Scoresbya are described from two quarries in the Middle Triassic Nymboida Coal Measures of the Nymboida sub-Basin in north-eastern New South Wales. The new species Nilssonia dissita and Taeniopteris adunca are based on previously unpublished material from Queensland together with conspecific material from Nymboida. An additional four new species from Nymboida are described; Taeniopteris nymboidensis, Linguifolium parvum,
Gontriglossa ligulata and Scoresbya carsburgii.
Manuscript received 1 March 2010, accepted for publication 29 May 2010.
KEYWORDS: Middle Triassic flora, Nymboida Coal Measures, palaeobotany, simple fossil leaves.
INTRODUCTION
This is the eighth paper of a series describing the early-middle Triassic Nymboida flora. Part 1 of this series (Holmes 2000) described the Bryophyta and Sphenophyta, Part 2 (Holmes 2001) the filicophyta, Part 3 (Holmes 2003) fern-like foliage, Part 4 (Holmes and Anderson 2005a) the genus Dicroidium and its fertile organs Umkomasia and Pteruchus, Part 5 (Holmes and Anderson 2005b) the genera Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris, Part 6 (Holmes and Anderson 2007) the Ginkgophyta and Part 7 (Holmes and Anderson 2008) the Cycadophyta. In this paper the simple leaves in the genera Nillsonia, Taeniopteris, Linguifolium and Gontriglossa together with the enigmatic lobed leaf Scoresbya carsburgii are described.
A description of the Coal Mine and Reserve Quarries, the source localities of our described material
together with a summary of the geology of the Basin Creek Formation, the Nymboida Coal Measures and the Nymboida Sub-Basin were provided in Holmes (2000).
METHODS
The material described in this paper is based mainly on collections made by the senior author and his family from two then-active Nymboida quarries (Coal Mine Quarry and Reserve Quarry) over a period of forty years. The specimens noted in Flint and Gould (1975), Retallack (1977), Retallack et al (1977) and Webb 1980 were examined in the collections of the Australian Museum, Sydney, the Department of Geology and Geophysics of the University of New England, Armidale and the Queensland Museum, Brisbane..
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
The University of Queensland PhD thesis on “Aspects of Palaeontology of Triassic Continental Sediments in South-East Queensland” by J.A.Webb (1980) included the descriptive taxonomy of fossils of simple leaves, similar to those that form the subject of this paper. In addition to his own extensive field collections Webb also examined all available and relevant material in State and private collections. Descriptive taxonomy in the past has so often been based on very limited and often fragmentary material. From Webb’s extensive range of material it was possible to gain a better understanding of species boundaries through the natural range of variation occurring within the fossil populations. On the basis of floral similarities, the Esk Formation (Toogoolawah Group) of south-east Queensland and the Nymboida Coal Measures of north-east New South Wales were deposited contemporaneously in the Anisian-Ladinian (Flint and Gould 1977, Rigby 1977). Regrettably most of Webb’s research was never published. Because of its relevance to this paper, two new species presented below are based on his original descriptions and types with Webb acknowledged as the author. Taxonomically comparable Nymboida specimens are illustrated and listed as “Additional Material”.
Since the completion of the research by Webb (1980) new studies have been published on similar taxonomic groups from other Gondwana Triassic floras that are relevant to this paper. Retallack (1980) reviewed the Middle Triassic Tank Gully flora of New Zealand and proposed a new combination for Linguifolium tennison-woodsii; Artabe (1985) described six Taeniopteris species from Los Menucos Formation of Argentina; Anderson and Anderson (1989), in their taxonomic revision of the SouthA frican Molteno gymnosperms described and extensively illustrated nine species of Taeniopteris, five species of Linguifolium and three species of Gontriglossa; Gnaedinger and Herbst (1998) described three species of Taeniopteris and three species of Linguifolium from El Tranquilo Group of Argentina; Gnaedinger and Herbst (2004a) described ten species of Taeniopteris from northern Chile, using a statistical analysis of venation characters; Gnaedinger and Herbst (2004b) described one Linguifolium sp also from northern Chile and Herbst et al (2005) listed one Taeniopteris sp. and two Linguifolium spp from the Lake District of Chile.
The Nymboida specimens are preserved in mudstones, siltstones and sandstones as carbonaceous compressions or impressions in which the gross morphology is usually well-preserved. However spores and cuticles have been destroyed by a tectonic heating event during the Cretaceous Period (Russel
1994). Therefore our identification of taxa is based only on characters of gross morphology.
The exact stratigraphic horizon or detailed source of much of our Nymboida specimens is uncertain as most were collected from fallen blocks during quarry excavations. The Coal Mine Qua ry has not been active for some twenty years but the high working face, although now rather weathered, provides an excellent exposure of beds that demonstrate the palaeo- environmental conditions at the time of deposition and was described by Retallack (1977). In 2006 the Reserve Quarry was bulldozed into a featureless bowl — “for restoration and safety purposes” and the fossiliferous horizons are now hidden.
The Nymboida material described in this paper has been allocated AMF numbers and is housed in the palaeontology collections of the Australian Museum, Sydney.
DESCRIPTIVE TAXONOMY
Without supporting cuticular evidence and lack of affiliation with any fertile structures for a definite systematic placement, the leaves described below are regarded as form genera in Gymnospermae — sedis incertae. On the basis of preserved cuticle Nilssonia leaves with haplocheilic stomata have been placed in the Cycadales and leaves of taeniopterid morphology may belong in several groups from ferns to cycads. Anderson and Anderson (2003) placed their Molteno Taeniopteris species in the Pentoxylales based on affiliation evidence and similarly they placed Gontriglossa in the Gnetopsida. The affinities of Linguifolium remain uncertain although Retallack (1980) suggested an affiliation with the seeds Carpolithus mackayi. Scoresbya has been speculated as being a fern, a seed fern, a member of the Caytoniales (Taylor and Taylor 2009) or even a pro- angiosperm (Weber 1995).
Gymnospermae incertae sedis Genus Nilssonia Brongniart 1825
Type species Nilssonia brevis Brongniart 1825
Nilssonia is a form genus that includes simple linear to oblanceolate leaves to irregularly pinnate leaves. It has a worldwide distribution and ranges from the Triassic to the Cretaceous. The main gross distinguishing character of the leaves is the dorsal attachment of the lamina which completely covers
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
the mid vein. The appearance of this character is often an artefact of preservation, eg the fossil may be an impression of the upper or lower leaf surface or an internal or external cast or mould that often masks the form and place of attachment of the lateral veins to the midrib.
The venation pattern of leaves from Gondwana localities differs somewhat from that of species described from the northern hemisphere in the more common bifurcation of the lateral veins and their straight and parallel course to the margin. Similar simple leaves in which the lamina does not completely cover the mid vein and without preserved cuticle are placed in the form genus Taeniopteris. Where cuticle information is available, the haplocheilic stomata and trichomes indicate cycadalean affinities. No cuticle is preserved on the Nymboida material. Some specimens in our Nymboida collections can be placed in a previously unpublished species as described by Webb (1980). Note this species is attributed to Webb.
Nilssonia dissita J.A.Webb sp. nov. Figures 1A—C; 2A, B; 7A
Selected synonymy
1917 Taeniopteris crassinervis (Feistmantel) Walkom, p.38, Pl. 1, fig. 2.
1975 Nilssonia cf. princeps (Oldham and Morris) Seward; Flint and Gould, p.71.
1980 Nilssonia dissita Webb, p. 87, P1.11, figs 3, 6, 8, Text figs 18 c, d (Unpubl.)
Diagnosis
Large simple leaf 65-150 mm wide; midrib 2.5— 4 mm in width; lamina covers whole of mid-vein; secondary veins arise from the dorsal surface of a moderately wide central rib at fairly acute angle, then curve broadly to run at 80°—90° to margin; individual veins frequently bifurcate once, usually as they leave the central rib, occasionally fork a second time; density of venation 9-16 / 10 mm.
Description (revised to include new Nymboida material)
Leaves are simple, oblanceolate with undulate to entire margins and wavy to smooth surface, tapering to obtuse apex. Length from c. 200 to >300 mm, the leaf base is not known; width at mid lamina ranges from 60 —150 mm. Lamina is dorsally attached and completely covering the mid vein. Lateral veins diverging from a mid point above the mid vein at an angle of 50°-70°, arching to run at a high angle (70° — 90°) straight and parallel to the margin. Many
Proc. Linn. Soc. N.S.W., 131, 2010
veins bifurcate once, usually as they leave the central rib; a few subsequently fork a second time but never anastomose; veins coarse with a density 9-16 / 10 mm. Mid vein when exposed ranges in width from 1-4 mm.
Holotype GSQ F12897
Type Locality Geological Survey of Queensland Locality 1552, Esk Formation, Toogoolawah Group
Additional material
GSQ12898, Esk Fm. UNEF13443, AMF120989, AMF 130180, AMF 130181, AMF130182, AMF 130183, all from Coal Mine Quarry, Nymboida CM. Also the material listed by Webb (1980), mostly from the Esk Formation of Queensland.
Name derivation dissitus — Latin — distant, apart, referring to the widely spaced venation.
Discussion
Previous material from Nymboida (Flint and Gould, 1975) was recognised by Webb (1980) as questionably belonging to this species. From our new collections specimen AMF 130180 is a block showing two leaves (Fig. 2B), one almost complete, preserved in almost three dimensions in white sandstone. The lamina of the more complete leaf, in places, completely covers the mid vein as can be seen by the lateral veins appearing to adjoin in mid lamina. The incomplete specimens AMF130182 (Fig. 2A) and AMF130183 both show sections of a leaf with adjoining lateral vein bases over the mid vein. In other parts of these leaves and similarly in the full length of AMF130181 (Fig.1C) the mid vein is exposed as an artefact of preservation. These leaves are included in this species based on the form, course and density of their veins and there being no evidence that the veins were laterally attached to the margin of the mid vein.
Nilssonia moretonti Walkom 1928
Figure 8A Synonymy 1928 Nilssonia moretonii Walkom, p. 466, Pl. 25, WES 25 Bs Te
1980 Nilssonia moretonii Walkom; Webb, P1 10, figs 1, 4, 6, 7. 1989 Taeniopteris moretonii (Walkom) Anderson
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
and Anderson, comb. nov. p. 376, fig. 3; p.547,
figs 5, 6.
Description
A simple strap-shaped leaf with entire or slightly lobed margins; complete leaf unknown, from 30 — 110 mm wide; lamina covering whole of mid vein; lateral veins departing from a central line above the mid vein at an acute angle immediately arching then proceeding straight and parallel to the margin. Veins frequently fork on leaving the central rib and again soon after; density 20 — 35 / 10 mm.
Nymboida Material
Known only froma single specimen, AMF 130184 from Coal Mine Quarry, base and apex missing, vein density in lower portion of lamina 30 / 10 mm becoming denser distally, to 40 / 10 mm, straight and parallel at a high angle across lamina and curving slightly upwards to the margin.
Discussion
This leaf fragment is placed in N. moretonii on the basis of the very dense venation and its mid dorsal attachment to the mid vein.
Anderson and Anderson (1989) transferred Nilssonia moretonii to the genus Taeniopteris without additional comment. Under “Intergeneric comparisons” those authors noted that entire specimens of Ni/ssonia can hardly be effectively distinguished from Taeniopteris and did not use the genus Nilssonia. Many of the leaves placed in Taeniopteris (see below) show evidence of lateral attachment of the lamina but towards the dorsal edge of the mid vein. The degree of the lamina overtopping of the mid vein makes for a subjective differentiation between Ni/ssonia and Taeniopteris in the absence of preserved cuticle.
Genus Taeniopteris Brogniart 1832
Type species Taeniopteris vittata Brongniart 1832
Taeniopteris is a form genus for simple strap- shaped leaves with entire lamina and occasionally forking lateral parallel venation running at a high angle to a prominent midrib and with unknown cuticle (Meyen 1987, Taylor and Taylor 1993, Anderson and Anderson 2003). Numerous species have been described world-wide from the Upper Carboniferous to Recent. While this leaf form is diverse and widespread it rarely occurs in abundance. Many
species have been erected for Gondwana Triassic material, often based on limited or dubious specimens that do little to demonstrate the natural variation within a species. Recent papers on Triassic South American TJaeniopteris have been useful but some species appear to be based on very few specimens (eg for Argentina, Artabe 1985, Gnaedinger and Herbst 1998. For material from Chile, Gnaedinger and Herbst (2004a) have used a statistical analysis of venation sequence for ten species of Taeniopteris. Triassic material from South Africa was described by DuToit (1927) and very comprehensive collections from the Molteno Formation by Anderson and Anderson (1989, 2003) who described ten species from 29 assemblages (localities) and used the “palaeodeme approach” and illustrated the range of variation in a species. From Australia there are numerous species in the literature but most have been based on fragmentary material, inadequate descriptions and have often been poorly illustrated. Rarely has the natural range of variation that may exist in a species been recognised. In our Nymboida collections taeniopterid leaves comprise c. 3% of numbered specimens. Few leaves, especially the larger forms, are found complete. Occasional bedding planes (possible sub—authocthonous assemblages) show numerous individual leaves resembling a natural autumnal-like leaf fall. In many specimens the leaf lamina appears to be dorsally attached to the midrib but without totally covering it as in Nilssonia.
In our Nymboida collections the majority of taeniopterid leaves fall within the range of variation as recognised by Webb (1980) from his examination of over 170 specimens, mostly from the Esk Formation for his unpublished species Zaeniopteris adunca which is here validated using his type specimen and slightly emended diagnosis. Other rare Nymboida leaves with clearly distinguishing characters are described as the new species 7: nymboidensis.
Sterile leaves of the enigmatic fern Ogmos adinus (Webb 1983, Holmes 2001) may be placed as a form species of Taeniopteris but are not included here.
Taeniopteris adunca J.A.Webb sp. nov. Figures 3A—H; 4A—C; 5A—C
Selected synonomy
1892 Taeniopteris sp. indet. Etheridge, p. 374, PI. 16, fig. 4.
1924 Taeniopteris (? Danaeopsis) crassinervis (Feistmantel) Walkom; Walkom, p. 84, Pl. 18, fiewSe
1925 Taeniopteris carruthersii, Tenison-Woods; Walkom, p. 85, text fig. 3.
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
1965 Taeniopteris aff. lentriculiforme (Etheridge) Walkom; Hill et al., PL. T8, Fig. 4.
1975 Taeniopteris aff. lentriculiforme (Etheridge) Walkom; Flint and Gould, PI. 3, figs 8, 9. 1980 Taeniopteris adunca sp. nov. Webb (unpubl.),
Pl. 23, figs 1-11; text figs 51 a-.
Diagnosis
Strap-shaped leaves, very variable in width; leaf surface rarely undulate; secondary veins always leave midrib at moderately acute angle, then quickly arch away and travel straight and parallel to the margin at 70°—90°; individual veins frequently bifurcate twice but anastomose very rarely; vein density ranging from 15 to 25 per 10 mm near the margin.
Description
Leaves elongate, strap-shaped; tapering gradually and fairly uniformly to a stout petiolate base and distally to an obtuse to acute rounded apex; very _ variable in size, from 9—60 mm in width and from 110 mm to >250 mm in length; lamina rarely undulate, margins entire. Midrib sometimes striate, appearing as a prominent groove or ridge, 1-2 mm wide in mid leaf and expanding basally to c. 3 mm. Leaf lamina attached to the dorsal edge of the mid vein without overlapping the dorsal surface. Lateral veins always leave the mid vein at a moderately acute angle (usually less than 45°) and arch rapidly within | to 2 mm then proceed straight and parallel to the margin at an angle of c. 75° — 85° and more acutely towards the apex. Veins fork close to the mid vein and then once or rarely twice across the lamina. Conjoining of the veins is rare. Density of the veins varies between populations and leaf sizes and averages c. 15—25 /10 mm near the margin.
Holotype UQF 18836
Type locality G. R. 380 551 Blackbutt 1: 63 360 Sheet, Esk Formation, Toogoolawah Group, Anisian—Ladinian
Illustrated specimens from Queensland
UQF18836, UQF72601, UQF18830, UQF2103, UQF72814, UQF72813, UQF72811, UQF21494, see ign:
Additional material
AMF 130185, AMF130186, AMF130187, AMF- 130188, AMF130189, AMF130190, AMF130191, AMF130193, AMF130194, AMF130215. All from Coal Mine Quarry, Nymboida CM.
Proc. Linn. Soc. N.S.W., 131, 2010
Name derivation
aduncus, Latin, bent inward, hooked, referring to the abrupt curvature of the lateral veins as they leave the midrib.
Discussion
Based on the detailed study of extensive collections of fossil plant material mainly from Queensland, J.A.Webb (1980, unpublished) differentiated two commonly occurring strap-like Taeniopteris leaf forms mainly on the basis of the form of attachment of the lateral veins to the mid vein. Taeniopteris carruthersii, widespread in the Upper Triassic assemblages, has lateral veins arising straight from the midrib at a high angle, sometimes forking and running at almost right angles to the leaf margin. In 7’ adunca the leaf lamina is attached dorsally to the midrib with the lateral veins diverging from the mid vein at an acute angle, usually forking close to the base then arching and running straight to the margin at a high angle. This arching of the veins close to the mid vein is often obscured through the form of preservation during fossilization but can be revealed from close examination. While there are wide variations within the two species and some overlapping characters, Webb recognised the two species as distinct and with stratigraphic implications. T. carruthersii occurs in the Late Triassic Ipswich Coal Measures whereas 7’ adunca is found in the Esk Formation of Queensland and the Basin Creek Formation of the Nymboida Coal Measures, both Middle Triassic units.
T. adunca is the most commonly occurring form of Taeniopteris at Nymboida. On some bedding planes (see blocks AMF130190, AMF130216, AMF130193 and AMF130194) the leaves form an almost mono- specific assemblage, probably a seasonal leaf-fall. Both within and between these assemblages there is a wide variation in leaf size and shape. 7: adunca is regarded as a species complex.
Taeniopteris parvilocus Anderson and Anderson from South Africa (Anderson and Anderson 1989) and from Chile (Herbst et al. 2005) is similar to T. adunca in outline and size but differs by the less dense venation (13/10 mm) that runs almost straight from the midrib and then arches upwards towards the margin. See below for comparisons with 7: nymboidensis.
Taeniopteris nymboidensis Holmes and Anderson sp. nov. Figures 6 A, B
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Diagnosis
Leaf oblanceolate, to 150 mm long, 30 mm wide; apex obtuse; lateral veins dorsally attached at acute angle to strong mid vein, widely spaced at point of attachment, c. 6/10 mm, arching through half the width of the lamina and then running straight to margin at c. 65°—70°, bifurcating in an irregular pattern, once near the base and again across the lamina; vein density in mid lamina c. 14—18/10 mm.
Description
Leaves simple, entire, oblanceolate to 150 mm long and from 25—30 mm wide, apex obtuse; strong mid vein 2 mm wide at mid lamina and tapering distally; base petiolate to >15 mm long. Lateral veins attached on dorsal edge of the mid vein, decurrent, widely spaced at point of attachment, c. 6/10 mm, arching then running straight and parallel to the margin at c. 65°—70° in mid lamina but more acute towards the base and apex. Most veins bifurcate while arching from the base and usually once again at irregular distances from the margin. The pattern of bifurcation is very irregular. Vein density in the mid lamina c. 14—18/10 mm.
Holotype AMF 130197
Type locality Coal Mine Quarry, Nymboida, Basin Creek Formation, Nymboida Coal Measures.
Other material AMF 130198, Coal Mine Quarry.
Name derivation nymboidensis- with reference to the type locality
Discussion
Only two slabs in the collections display this new species. The holotype is on a block on which are the remains of seven leaves, four appearing to arise from a common point but the point of attachment is not preserved (Fig. 6A). 7: nymboidensis differs from T adunca by its oblanceolate shape, by the arching of the lateral veins which continues half way across the lamina and by the irregular bifurcation of the lateral veins. In shape and venation pattern 7. nymboidensis is similar to 7: troncosoi Gnaedinger and Herbst (2004a) but differs by the less dense venation. 7° fissiformis Anderson and Anderson (1989) is similar to T. nymboidensis in vein density (15/10 mm) but is a much smaller leaf; 77 anavolans Anderson and Anderson (1989) is similar in shape and size but has coarser venation of c. 12/10 mm.
Taeniopteris sp A Figure 7B
Description
Mid portion of a very large leaf >100 mm wide; mid-vein to 5 mm wide, longitudinally striate; lateral veins attached to the dorsal edge of the mid vein at 60°—70° and quickly arch and run at c. 80° straight and parallel to each other across the lamina and curve slightly upwards towards the margin. Some of the lateral veins bifurcate close to the mid-vein and others occasionally fork at varying distances towards the margin. The vein density is ca 10—12/10 mm.
Material AMF 130199 Coal Mine Quarry.
Discussion
This fragment differs from 7? adunca and T. nymboidensis by the larger size and broader mid vein and from N. dissita by the lateral veins not overtopping the mid vein. Zaeniopteris sp. A of Anderson and Anderson (1989) from the Triassic Molteno Formation of South Africa is a very much larger leaf with a finer mid rib and lateral veins almost overtopping the mid vein. Another large leaf from the Molteno Formation, Taeniopteris homerifolius Anderson and Anderson (1989) has a venation pattern with veins upcurving towards the margin similar to 7: sp. A but differs by the lateral attachment of the lamina to the mid- vein. Webb (1980 p. 218) described a Taeniopteris sp. (unpublished) with much larger leaves — to 240 mm wide and lateral veins occasionally anastomosing which he compared with a leaf from South Africa described by DuToit (1927) as Taeniopteris lata.
Genus Linguifolium Arber 1913 emend. Retallack 1980
Type species Linguifolium lilleanum Arber 1913
Linguifolium was erected for simple entire leaves, linear, spathulate, lanceolate or obovate; apices sub-acute to rounded; with mid vein persistent to apex; lateral veins arising at very acute angle to the mid rib then arching to meet the margin at an acute angle, forking once and occasionally twice in the nearer third of their length. The status of the genus Linguifolium was well-discussed by Retallack (1980). Linguifolium leaves are extremely rare in the Nymboida collections.
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Linguifolium tennison-woodsii (Jack and Etheridge 1892) Retallack 1980 Figures 8B, C
Selected synonymy
1892 Angiopteridium tennison-woodsii, Jack and Etheridge, p. 365
1898 Taeniopteris tennison-woodsii, Shirley, comb. nov. p. 23, Pl. 9, fig. 2.
1947 Doratophyllum tennison-woodsii, Jones and deJersey, p.37, Pl. 6, fig. 1.
1980 Linguifolium tennison-woodsii, Retallack, comb nov. fig. 7 F—H.
1980 Linguifolium tennison-woodsii, Webb, p.172, Pl. 20, figs 1-4, P1.21, figs 1-15, text fig. 41, a—p, (unpubl.).
1989 Linguifolium tennison-woodsii, Anderson and Anderson, p.522, figs 1-3.
1998 Linguifolium tennison-woodsii, Gnaedinger and Herbst, P1.1, fig. d.
Description
A portion of a small linear leaf with the base missing, tapering slightly distally to an incomplete apex. Length preserved 80 mm, width 6 mm. Mid vein not well defined, lateral veins decurrent on mid vein, arching across lamina to meet entire margin at c. 75°, forking once close to mid vein. Vein density in mid lamina c.12/10 mm.
Material AMF 130200, Coal Mine Quarry, Basin Creek Formation, Nymboida Coal Measures.
Discussion
Linguifolium tennison-woodsii differs from most Linguifolium spp. by its narrow linear form and from the extremely narrow Linguifolium gracile from the Molteno of South Africa (Anderson and Anderson 1989) by its more arching and denser veins.
Linguifolium parvum sp. nov. Holmes and Anderson 2010 Figures 9A—C
Diagnosis
Small spathulate sessile leaves less than 100 mm long, lateral veins decurrent on striated mid vein, arching across lamina to meet margin at acute angle, number of veins forking near base variable, very occasional veins forking and conjoining. Vein density 8-12/10 mm.
Proc. Linn. Soc. N.S.W., 131, 2010
Description
Leaf spathulate; maximum length 100 mm; width from 11—20 mm, apex rounded, lamina tapering to sessile base; midrib with longitudinal striations, width at base 1.5 mm, contracting in width through length of the leaf; lateral veins decurrent, arching from mid- vein across lamina to reach the margin at an angle of 30°-45°; c. half the veins fork once close to the mid vein; occasional veins fork in the mid lamina and conjoin to form a long narrow areole. Density of the veins at mid lamina ranges from 8 to 12/10 mm.
Holotype AMF130201
Type locality Coal Mine Quarry, Basin Creek Formation, Nymboida Coal Measures.
Other Material
AMF 130202, AMF130203, AMF130204, and AMF 130207 from Coal Mine Quarry. AMF 130205 and AMF 130206 from Reserve Quarry.
Name derivation parvum — Latin — small, referring to the small size of the leaves of this taxon..
Discussion
Linguifolium parvum is similar in form to L. lilleanum Arber (1913), L. ascium Webb (1980) and L. patagonicum Gnaedinger and Herbst (1998) but differs by the short length and by the density and course of the lateral veins. In the Nymboida collections these Linguifolium leaves are very rare. The generic diagnosis of Linguifolium states that the lateral veins do not anastomose. However on some specimens of L. parvum very occasional lateral veins fork and conjoin to form a long narrow areole, hardly reason to remove it from Linguifolium.
? Linguifolium sp. A Figures 8D, E
Description
A small spathulate leaf somewhat resembling in shape L. parvum, is 74 mm long and 14 mm wide, with base and apex missing. The lateral veins are sparse, c. 8/10 mm and arch slightly across the lamina at c. 45° to each terminate at a tooth along a unique finely serrate margin; occasional veins forking once between mid vein and mid lamina.
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Material AMF 130208 and counterpart AMF 130209, Coal Mine Quarry.
Discussion
This form is based on a single specimen and its counterpart. It differs from all described species of Linguifolium by the serrate margin. Jungites polymorpha from the Molteno Formation (Anderson and Anderson 1989) has a finely serrate margin but differs by the dense parallel venation and the variably entire to pinnate lamina margin.
Genus Gontriglossa Anderson and Anderson 1989
Type species Gontriglossa verticillata (Thomas 1958) Anderson and Anderson 1989
The genus Gontriglossa was erected by Anderson and Anderson (1989) for elliptic, petiolate leaves with veins attached at an acute angle, arching and anastomosing towards the margin. Some specimens of G. verticillata from the Molteno Formation of South Africa (Anderson and Anderson 1989, 2003) show stems with well-spaced opposite fascicles of three leaves. From Nymboida, Holmes (1992) described some reticulate veined leaves that were identified as Triassic “G/ossopteris-like leaves”. Those leaves are here transferred to the genus Gontriglossa. Amongst the Nymboida material is a specimen showing 10 leaves attached in a whorl or a close spiral (10A, 12A). To accommodate this form in Gontriglossa requires a slight emendation of the generic diagnosis to include the attachment of leaves as either terminal whorls, close spirals or well-spaced opposite fascicles.
Gontriglossa grandis (Walkom) Holmes and Anderson comb. nov. Figures 10A; 12A
Synonymy
1928 Anthrophyopsis grandis Walkom, p. 464, text fio: 2, Pls 265 fie. S:
1992 ?Glossopteris grandis Holmes, p. 122, Pl. 2, figs1, 2.
Description
Leaves oblanceolate, to 150 mm long, and to 95 mm wide but usually much smaller, attached as a terminal whorl or a close spiral, apex rounded acute to obtuse, tapering basally to a short petiole; midrib
distinct, striate; lateral veins leave the midrib at an acute angle and for about one third of the width of the lamina they bifurcate and anastomose to form a wide elongate mesh with a general inclination of c. 45° to the midrib; for the remainder of the lamina they form a narrower elongate mesh inclined at 65°—70° to the midrib; closer to the midrib the meshes are 1-2 mm wide, wider in the proximal than the distal part, while towards the margin they narrow to form 7-8 meshes per 5 mm of width.
Holotype UQF1724-5, University of Queensland, Brisbane from Sheep Station Creek in the Esk Beds.
Other material AMF 78254-78258, Australian Museum, Sydney — from Coal Mine Quarry, Nymboida.
Discussion
The Nymboida leaves placed in this species are much smaller (c. 80 mm long and c. 30 mm wide) than the holotype specimen but are closely similar in gross form and the anastomosing venation pattern. The Nymboida specimens are notable for the whorled or closely spiral arrangement of the leaves. Individual leaves of G. verticillata (Thomas) Anderson and Anderson (2003) are similar in size and venation pattern to the Nymboida leaves but differ by the known cuticle and the well-spaced opposite attachment of fascicles of three leaves to an elongated stem.
Gontriglossa nymboidensis Holmes and Anderson comb. nov. Figures 11A, B
Selected Synonymy .
1975 Anthrophyopsis grandis Walkom, Flint and Gould, Pl. 1, fig. 9.
1992 ?Glossopteris nvmboidensis Holmes, P. 122, Pl. 1p esiSt4 Ply 2ifieele
Holotype
UNEF13528 and paratype UNEF13639, both from Coal Mine Quarry. Now housed in the Australian Museum as specimens AMF 126731 and AMF 126730 respectively.
Additional material AMF 130214, Coal Mine Quarry.
Description A reticulate veined leaf known only from apical
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W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
and mid lamina fragments. Leaf of unknown length, width 50 mm, tapering distally to an acutely rounded apex; midrib distinct, striated; lateral veins leaving midrib at c. 20°-30° at intervals of ca 0.5 mm and quickly arch over a distance of c. 5 mm where they bifurcate and then run straight to the margin at an angle of 75°. After the initial bifurcation the veins fork again two or three times to join with adjacent veins to form long narrow meshes, each subsequent mesh being narrower than the proceeding one. The density of the veins in the mid lamina is c. 12—14/ 10 mm and at the margin c. 18/10 mm.
Additional material AMF130214, Coal Mine Quarry.
Discussion
G. nymboidensis differs from all other Gontriglossa species by the very fine narrow parallel meshes formed by the lateral veins. Cetiglossa balaena _ Anderson and Anderson (2003) from the Molteno of South Africa is much larger leaf with more elongate reticulate venation that does not arch from the mid vein. The somewhat similar reticulate veined leaf from Patagonia, Santacruzia hunickenii Gnaedinger and Herbst (1998) differs by the serrate to incised margins and the lateral veins attached at a high angle and running straight to the margin. (See comparison of Santacruzia hunickenii with Gontriglossa lacerata below).
Gontriglossa lacerata (Holmes 1992) Holmes and Anderson comb. nov. Figures 11C, D
Synonymy 1992 ?Glossopteris lacerata Holmes, p. 124, Pl. 2,4.
Holotype AMF78259. Coal Mine Quarry, Basin Creek Formation, Nymboida Coal Measures.
Additional material AMEF130210 and AMF130213 from Reserve
Quarry
Description
Known from three incomplete specimens. Leaf broad-elliptic or oblanceolate, >180 mm long, 65 mm wide, petiolate; apex broadly rounded; margin iregularly lacerate, dentate or lobed; venation somewhat similar to G. nymboidensis, arching from mid-vein, bifurcating and anastomosing to the margin.
Proc. Linn. Soc. N.S.W., 131, 2010
Discussion
This is a bizarre species. It differs from other Gontriglossa species by the irregularly lacerate margins which we believe to be natural and not resulting from insect damage.
Gnaedinger and Herbst (1998) described from the Triassic Tranquilo Group of Santa Cruz, Argentina a leaf with reticulate venation and serrate to deeply incised margins and placed it in their new genus and species Santacruzia hunickenii. They were perhaps unaware of the paper by Holmes (1992) as they made no comparisons with ?G/ossopteris (now Gontriglossa) lacerata. S. hunickenii differs from Gontriglossa retculata by the less deeply incised margin and by the much denser venation that passes at 90° from the mid-vein to the margin. Gnaedinger and Herbst did compare Santacruzia with the Molteno species Gontriglossa balaena that has been transferred to the genus Cefiglossa Anderson and Anderson (2003) which lacks the lacerate lamina margin.
Gontriglossa ligulata Holmes and Anderson sp. nov. Figures 12B—D
Diagnosis
Leaf ligulate, lateral veins decurrent on mid vein, widely spaced, arching and bifurcating once then running straight at a high angle towards the margin; forking again in mid lamina and conjoining to form a longitudinal row of transverse rhomboidal areoles and a row of triangular areoles parallel and adjacent to the margin.
Description
An incomplete strap-shaped leaf 80 mm long but with base and apex missing; lamina 14 mm wide above broken base, tapering gradually over whole length to 8 mm; mid vein 1 mm wide; lateral veins decurrent and widely spaced on mid vein, arching and bifurcating once then passing to margin at c. 75°. Between mid lamina and margin each vein bifurcates twice and anastomoses with adjacent veins to form a longitudinal row of transverse rhomboidal areoles and a row of triangular areoles parallel to the margin; vein density near margin c. 16/10 mm.
Holotype AMF 130211
Type Locality Reserve Quarry, Nymboida, Basin Creek Formation, Nymboida Coal Measures.
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Name derivation ligulata — Latin, strap-shaped, referring to the broad-linear form of the leaf.
Discussion.
This new species is based on a single incomplete specimen. While recognising that some species of Taeniopteris, eg T. fissiformis and T: anavolans (Anderson and Anderson 1989; Gnaedinger and Herbst 2004a) may show rare and irregular anastomoses, we believe that from the regular and distinctive anastomosing venation (see Fig. 12D) this leaf is best placed in Gontriglossa, The linear shape of the leaf and the details of the anastomosing venation pattern differentiate G. /igulata from the other Gontriglossa species described above and from the cordate based leaf, G. hilaryjanea (Anderson and Anderson 1989, 2003). The regular form of the marginal areoles diffentiates G. /igulata from the Scoresbya sp. described below.
Genus Scoresbya Harris 1932
Type species Scoresbya dentata Harris 1932
Scoresbya dentata was described by Harris (1932) for small palmate leaves with reticulate venation and dentate margins from Scoresby Sound in the Jurassic of Greenland. Additional specimens of Scoresbya dentata have been described from the Jurassic of Germany (Krausel and Schaarschmidt 1968), from China (Cao 1982), Afghanistan and Iran (Schweitzer and Kirchner 1998) plus an additional species from the Late Triassic of Mexico (Weber 1995). An incomplete specimen showing parts of several segments of a palmate leaf with dentate margin and reticulate venation from the Ipswich Coal Measures of Queensland was described by Shirley (1898) as Phlebopteris (?) dichotoma and later transferred by Herbst (1974) to the Scoresbya genus.
Scoresbya carsburgii Holmes and Anderson sp. nov. Figures 13A, 14A, B.
Diagnosis
A large leaf bifurcating irregularly into broad linear lobes; margins entire to irregularly serrulate; lateral veins decurrent on striate mid vein, then arching and running to margin, forking near base, occasionally in mid lamina and then forking and
10
sometimes conjoining to form small areoles adjacent to the margin; vein density in mid lamina c. 12 / 10 and c. 18 / 10 mm near margin.
Description
An incomplete palmate leaf; mid _ vein longitudinally striated, 3 mm wide in proximal section of leaf; lamina bifurcating at 10 mm from the base of leaf as preserved. The minor fork produces a broad linear pinna or lobe 90 mm long and 28 mm wide. After 43 mm the main rachis again bifurcates to form a major elongate lobe (pinna) 120 mm long and 30 mm wide and a minor lobe 60 mm long and 20 mm wide, both tapering slightly distally. The margins of the lobes are entire to irregularly undulate or serrulate. Throughout the leaf the decurrent lateral veins are widely spaced as they arch at an acute angle from the main rachis, soon forking irregularly and then running straight to the margin at c. 30°-45°, again sometimes forking at irregular distances across the lamina; close to the margin some veins again fork and conjoin to form small triangular areoles adjacent and parallel to the margin (Fig. 14B). Density of the lateral veins in mid lamina c 12/10 mm and near the margin c 18/10 mm.
Holotype AMF130212
Type Locality Reserve Quarry, Nymboida, Basin Creek Formation, Nymboida Coal Measures.
Name derivation
carsburgii — named for the collector of the specimen, amateur fossil plant and insect enthusiast, Mr Allan Carsburg.
Discussion
Scoresbya carsburgii is based on a single incomplete specimen that overlies another lobe fragment. It differs from the northern hemisphere species S. dentata Harris by its larger size, less obvious dentate or pinnatifid margins and by the form of venation. Scoresbya dichotoma (Shirley) Herbst (1974) from the Ipswich Coal Measures of Queensland is a smaller leaf and as described by Herbst has veins conjoining to form an intramarginal vein similar to that in the genus Yabiella. From the late Triassic of Chile Mollesia melandeziae Melchior and Herbst (2000) is described as particularly similar to Scoresbya but with a different venation pattern. The affinities of Scoresbya are not well understood. Herbst (1992) excluded Scoresbya from
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
the Dipteridaceae and Taylor et al (2009) discussed it under the Caytoniales while Weber (1995) inferred a possible link with angiosperms. S. carsburgii is an interesting addition to the Nymboida flora and illustrates the many puzzles still to be solved in these ancient floras.
CONCLUSION
This paper deals with leaves of simple form placed in the form genera Nilssonia, Taeniopteris, Linguifolium and Gontriglossa and a unique lobed leaf referred to the genus Scoresbya. Described are
two species of Nilssonia including a new species N.
dissita; three species of Taeniopteris including the new species 7. adunca and T: nymboidensis; two species of Linguifolium including the new species L. parvum; four species of Gontriglossa including three new combinations and a new species G. /igulata. _ A unique specimen of a lobate leaf is described as Scoresbya carsburgii sp. nov.
ACKNOWLEDGEMENTS
WBKH deeply appreciates the assistance provided by his daughters Marnie and Netta and late wife Felicity in collecting from the Nymboida localities over many years. Drs Susan Parfrey and Kristen Spring of the Queensland Museum Collections kindly located specimens described in Webb’s Thesis. WBKH is assisted by a grant from the Betty Main Research Fund.
REFERENCES
Anderson, J.M and Anderson, H.M. (1989). Palaeofiora of southern Africa. Molteno Formation (Triassic). Vol.2: Gymnosperms (excluding Dicroidium). Balkema, Rotterdam.
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Artabe, A.E. (1985). Estudio systematico de la tafoflora Triasica de Los Menucos, provincial de Rio Negro, Argentina. Parte 2. Cycadophyta, Ginkgophyta y Coniferophyta. Ameghiniana 22, 159-180.
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Cao, Z.Y., 1982. On the occurrence of Scoresby from Jiangsu and Weichselia from Zhejiang. Acta Palaeontologica Sinica 21, 344-348.
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Gnaedinger, S. and Herbst, R., (2004a). Pteridophylla del Triasico del Norte Chico de Chile. 1. El género Taeniopteris Brongniart. Ameghiniana 41, 91-110.
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Triassic Fossils of Queensland. Queensland Palaeontographical Society, Brisbane. 1-32.
Holmes, W.B.K. (1992). Glossopteris-like leaves from the Triassic of eastern Australia. In: Venkatachala, B.S., Jain, K.P. and Awasthi, N. Eds. Proceedings of the ‘Birbal Sahni Centenary Palaeobotanical Conference’, Geophytology 22, 119-125.
Holmes, W.B.K. (2000). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 1. Bryophyta, Sphenophyta. Proceedings of the Linnean Society of NSW 122, 43-68.
Holmes, W.B.K. (2001). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 2. Filicophyta. Proceedings of the Linnean Society of NSW 123, 39-87.
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Holmes, W.B.K. (2003). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 3. Fern-like foliage. Proceedings of the Linnean Society of NSW 124, 53-108.
Holmes, W.B.K. and Anderson, H.M. (2005Sa). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 4. Dicroidium. Proceedings of the Linnean Society of NSW 126, 1-37.
Holmes, W.B.K. and Anderson, H.M. (2005b). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 5. The Genera Lepidopteris, Kurtziana, Rochipteris and Walkomiopteris. Proceedings of the Linnean Society of NSW 126, 39-79.
Holmes, W.B.K. and Anderson, H.M. (2007). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 6. Ginkgophyta. Proceedings of the Linnean Society of NSW 128, 155-200.
Holmes, W.B.K. and Anderson, H.M., (2008). The Middle Triassic flora of the Basin Creek Formation, Nymboida Coal Measures, New South Wales. Part 8. Cycadophyta. Proceedings of the Linnean Society of NSW 129, 113-140.
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Melchior R.N. and Herbst, R., (2000). Sedimentology of the El Puquén Formation (Upper Triassic, Central Chile) and the new plant Mollesia melendeziae gen. et sp. nov. (pteridophylla, incertae sedis). Ameghiniana 37, 477-485.
Meyen, S.V., (1987). Fundamentals of Palaeobotany. Chapman and Hall, New York.
Ottone, E.G. (2006). Plantas triasicas del Grupo Rincon Blanco, Provincia de San Juan, Argentina. Ameghiniana 43, 477-486.
Retallack, G.J. (1977). Reconstructing Triassic vegetation of eastern Australia: a new approach for the biostratigraphy of Gondwanaland. Alcheringa 1, 247-278. Alcheringa-fiche 1, G1—J16.
Retallack, G.J., Gould, R.E. and Runnegar, B. (1977). Isotopic dating of a middle Triassic megafossil flora from near Nymboida, north-eastern New South Wales. Proceedings of the Linnean Society of NSW 101, 77-113.
Retallack, G.J., (1980). Middle Triassic megafossil plants and trace fossils from Tank Gully, Canterbury,
New Zealand. Journal of the Royal Society of New Zealand. 10, 31-63.
Rigby, J.F. (1977). New collections of plants from the Esk Formation, south-eastern Queensland. Queensland Government Mining Journal 78, 320-325.
Russel, N.J., (1994). A palaeothermal study of the Clarence-Moreton Basin. Australian Geological Survey Organisation Bulletin 241, 237-276.
Schweitzer, H.J. and Kirchner, M., 1998. Die rhato- jurassischen Floren des Iran, Afghanistan. 11. Ptreidophyta und Cycadophyta. 1. Cycadales. Palaeontographica 248B, 1-85
Shirley, J. (1898). Additions to the fossil flora of Queensland. Queensland Geological Survey Bulletin 7, 19-25.
Taylor, T.N. and Taylor, E.L. (1993). The biology and evolution of fossil plants. Prentice Hall, New Jersey.
Taylor, T.N., Taylor, E.L. and Krings, M., (2009) Palaeobotany: The Biology and Evolution of Fossil Plants. Academic Press. Burlington MA.
Walkom, A.B. (1917). Mesozoic floras of Queensland. Part | (contd.) The flora of the Ipswich and Walloon Series. (d) Ginkgoales, (e) Cycadophyta, (f) Coniferales. Queensland Geological Survey Publications 259, 1-49
Walkom A.B. (1924). On fossil plants from Bellevue, near Esk. Memoirs of the Queensland Museum 8, 77-92.
Walkom A.B. (1925). Notes on some Tasmanian Mesozoic plants.Part 1. Papers and Proceedings of the Royal Society f Tasmania 1924, 73-89.
Walkom A.B. (1928). Fossil plants from the Esk district, Queensland. Proceedings of the Linnean Society of NSW 53, 458-468.
Webb, J.A. (1980). Aspects of the palaeontology of Triassic continental sediments in South-East Queensland. Unpublished Thesis. Geology Department, University of Queensland.
Webb, J.A., 1983. A new plant genus, possibly a Marattealean fern from the Middle Triassic of eastern Australia. Memoir of the Association of Australasian Palaeontologists 1, 363-371.
Weber, R., 1995. A new species of Scoresbya Harris and Sonoraphyllum gen. nov. (Plantae incertae sedis) from the Late Triassic of Sonora, Mexico. Revista Mexicana de Ciencias Geologicas 12, 94—107.
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Figure 1. A-C. Nilssonia dissita Webb sp. nov. A. GSQF12897, Holotype, GSQ Locality 1552, Esk Fm. B. GSQF12898, GSQ Locality 1552, Esk Fm. C. AMF130181 Coal Mine Quarry, Nymboida CM. Scale bar = 1 cm.
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PRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
14
Figure 2. A. B. Nilssonia dissita Webb sp. nov. A. AMF130182, Coal Mine Quarry. Scale bar = 5 cm. B. AMF130180, Coal Mine Quarry, Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 3. A-H. Taeniopteris adunca Webb sp. nov. A. UQF18836, Holotype. 380 551 Blackbutt Sheet. B. UQF72601, UQL4110. C. UQF18830, 445 486 Blackbutt Sheet. D. UQF2103. UQL4238. E. UQF72814, UQL4255. F. UQF72813, UQL4238. G. UQF72811, UQL4110. H. UQF21494, UQLS585. All from Esk Fm. Scale bar = 1 cm
Proc. Linn. Soc. N.S.W., 131, 2010
15
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Figure 4. A-C. Taeniopteris adunca Webb sp. nov. AMF 130194, Reserve Quarry. B. AMF130195, Coal Mine Quarry. C. AMF130186, Coal Mine Quarry. All Nymboida CM. Scale bar A, C = 1 cm, B =S'cmi:
16 Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 5. A— C. Taeniopteris adunca Webb sp. nov. A. AMF130187. B. AMF130189. C. AMF130196, all from Coal Mine Quarry. Nymboida CM. Scale bar A, B= 1 cm. C =5 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
17
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Figure 6. A, B. Taeniopteris nymboidensis Holmes and Anderson sp. nov. A. AMF130197. B. AMF130198, both from Coal Mine Quarry. Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 7. A, B. Nilssonia dissita Webb sp. noy. AMF120939. B.Taeniopteris sp A. AMF 130199, both Coal Mine Quarry. Nymboida CM. Scale bar = 1 cm.
Roe. Linn Soe N.S WwW. als 1, 200
19
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
Figure 8. A. Nilssonia moretonii AMF130184. B, C. Linguifolium tennison-woodsii AMF130200. D, E. Linguifolium sp A AMF130208. Numboida CM. Scale bar A, C, E = 1 em, B=5 cm.
20 Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 9. A— E. Linguifolium parvum Holmes and Anderson sp. nov. A, B. Holotype AMF130201, Coal Mine Quarry. C, D. AMF130207, Coal Mine Quarry. E,. AMF130206, Reserve Quarry. Nym-
boida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
21
TRIASSIC GYMNOSPERMAE FROM NY MBOIDA - SEDIS INCERTAE
Nw NM
Figure 10. A. Gontriglossa grandis (Walkom) Holmes and Anderson comb. nov. Holotype AMF 78254 Coal Mine Quarry. Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 11. A, B. Gontriglossa nymboidensis (Holmes) Holmes and Anderson comb. nov. A. Holotype AMF126730. Coal Mine Quarry. B. Paratype AMF126731. Coal Mine Quarry. C, D. Gontriglossa lacerata (Holmes) Holmes and Anderson comb. nov. C. Holotype AMF78259 Coal Mine Quarry.. D. AMF130210, Reserve Quarry. Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010 ™®
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
24
Figure 12. A. Gontriglossa grandis (Walkom) Holmes and Anderson comb. nov. AMF78254 Coal Mine Quarry. B —D. Gontriglossa ligulata Holmes and Anderson sp. nov. AMF130211, Reserve Quarry. Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
W.B.K. HOLMES, H.M. ANDERSON AND J.A. WEBB
Figure 13. A. Scoresbya carsburgii Holmes and Anderson sp. nov. Holotype AMF130212, Reserve Quarry. Nymboida CM. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
DS
TRIASSIC GYMNOSPERMAE FROM NYMBOIDA - SEDIS INCERTAE
26
Figure 14. A, B. Scoresbya carsburgii Holmes and Anderson sp. nov. A. Line drawing of Holotype. AMF130212. B. Details of venation. Scale bar = 1 cm.
Proc. Linn. Soc. N.S.W., 131, 2010
Catalogue of Insects Collected by William Sharp Macleay in Cuba 1825-1836
Dominic Cross!. AND ELIZABETH JEFFERYS! .
'The University of Sydney, Faculty of Agriculture, Food and Natural Resource, NSW 2006 (dero3102@uni.sydney.edu.au) *EAJ Consultants” Principal, 14 Holloway Street, Pagewood NSW 2035 (liz@eaj.com.au)
Cross, D. and Jefferys, E. (2010). Catalogue of insects collected by William Sharp Macleay in Cuba 1825-1836, Proceedings of the Linnean Society of New South Wales 131, 27-35.
All of William Sharp Macleay’s labelled Cuban insects are now in a separately labelled Cuban insect cabinet in the Macleay Museum. There are over 7,349 labelled, pinned and partially identified. Other unlabelled specimens are still to be found throughout the collection. The geographical area where Cuba lies is also within the bio-geographical area for the southern United States, the Bahamas, the Caribbean and the northern most areas of South America. The biological scientists of these surrounding countries will find the information and knowledge of the distributions of insects of Cuba found in 1825 to 1836 of tremendous interest in relation to the possible distributions of insect faunas found or no longer found in
these areas today.
Manuscript received | March 2010, accepted for publication 24 May 2010.
KEYWORDS: Catalogue, Coleoptera, Cuba, Cuban insects, Curculionidae, Havana, Hymenoptera, Lepidoptera, Macleay Museum, Slave trade, William Sharp Macleay.
INTRODUCTION
The following is a catalogue of Cuban insects collected by William Sharp Macleay during his appointment as commissioner for the abolishment of the slave trade in Havana from 1825 to 1836. The specimens were taken from Cuba to England at the conclusion of his posting and consequently were moved to Sydney, Australia, with W.S. Macleay when he moved there to live. The collection of over 7000 insects were spread throughout the Macleay Museum’s entomology collection but were readily identified using locality labels. This is the first account of the Macleay Cuban collection and although initially the collection may have been larger, it is probable that over 170 years, specimens have had labels removed, been damaged beyond usefulness, or removed from the Macleay Museum altogether. All the remaining labelled Cuban specimens are now reunited in a single collection and are for the most part in good condition.
The collection consists of 7349 insects across at least 11 orders as follows:
Blattodea 33 Coleoptera 2172 Diptera 385 Hemiptera 12D Hymenoptera 3509 Lepidoptera 407 Neuroptera 40 Odonata 24 Orthoptera ] Phasmatodea 20 Siphonaptera 29
While care was taken to provide the most up to date species names, information was not able to be found on some of the labelled species name, and these have been included as written on the label. Where the year has been omitted it is where we were unable to find the complete documentation of the description and the publication.
William Sharp Macleay left England for Cuba in October 1825, to take up his duties in connection with the Mixed British and Spanish Court of Commission for the Abolition of the Slave Trade established at
THE MACLEAY COLLECTION OF CUBAN BEETLES
Havana. His residence in Cuba lasted from December 1825 to early in the year 1836. The catalogue of insects included in this paper, includes all those insects (over 7000) that are clearly labelled with the locality Cuba. William collected many specimens during those eleven years in Cuba, and then brought them to Australia. All of William Sharp’s collection is now housed in the Macleay Museum at the University of Sydney. There may be many more Cuban insects in the Macleay Museum but this catalogue only deals with specimens with the label Cuba.
William Sharp Macleay was born in London, on 21“ July 1792, the eldest son of Alexander Macleay (1767 - 1848) who amassed probably the finest insect collection in Europe and which eventually Alexander brought with him to Australia in 1826. William Sharp Macleay arrived in Australia in 1839 with his own insect collection from European collecting trips, his collection from Cuba and a collection of insects from his trip to the United States with Mr Titian Peale (Fletcher 1920).
William was educated at Westminster and Trinity College Cambridge and graduated with a BA in 1814 and MA in 1818. On leaving the University he was appointed as Attache to the British Embassy in France. What awakened and developed his interest in Zoology seems primarily to have been his father’s example, influence and fine collection of insects. During his time in Paris he had the opportunity of meeting Cuvier, Latreille and other distinguished naturalists of that time, as well as appreciating the importance of the magnificent establishment of the Jardin des Plantes. He subsequently was appointed Secretary to the Board for liquidating British claims on the French Government, established at the peace of 1815 - 1825. He was then sent as Commissioner of Arbitration of the Slave Trade established at Havana in Cuba. In 1830 he became the Commissary Judge of the same court. In 1836, he was appointed to be the Judge of the mixed British and Spanish Court of Justice, established under the treaty of 1835 — 1836. In 1836 he returned to England. In 1837 he retired from the Public Service. He left England in 1838 for Australia with his cousins William and John and arrived in Sydney in March 1839. Here he continued to collect insects and studied marine life. He was also a trustee of the Australian Museum from 1853 - 1862. He was universally recognised as the leading zoologist in Sydney from 1839 up to the time of his death. William Sharp died in Sydney on the 26" January 1865 and was buried in the family tomb in Camperdown Cemetery (Fletcher 1920).
William Sharp’s published work began in 1819 and ended in 1847 (over 30 published papers). There
28
were no publications on any of the insects that he collected in Cuba.
During his voyage to Cuba, in the months of October, November and December of 1825, he made notes on the Ornithology of the Islands of Madeira, Teneriffe and Saint Jago, as well as observations at Barbados, Martinique and off the coast of Saint Domingo. He always seemed to be taking notes of his natural surroundings wherever he went. However there seems to be no detailed notes of his insect collecting in Cuba, or at least none that has been found. However there is one interesting letter he wrote to his trusted friend Kirby, dated 3" 1827 January, about a year after his arrival. William writes:
“The climate has, I thank God, hitherto agreed with me much better than that of England: but there is a languor attendant upon every kind of exertion, which makes reading or study here a very different thing from what it is in England.” “This is a good place for Wading Birds, Lizards, Butterflies and Sphinges, (a term meaning Hawk Moths ), but apparently nothing else. I live in the country, where I have a large house and garden: this is my principal amusement, as I take great pleasure in cultivating Orchideae, particularly those which are parasitical on trees. The disagreeable are ants, scorpions, mygales and mosquitoes. The latter were quite a pest on my first arrival within the tropics, but now I mind them as much as I did gnats in England. “
The place of his residence in Cuba was Guanabacoa, (an Indian name meaning “site of the waters “) which he described as if “ living in the country is a picturesquely situated amid woods, on high hills which furnish a fine view, is a town a few kilometres from the capitol of Cuba, Havana.”
During his leisure hours, natural history soon began to claim his attention as he sent specimens of lizards, bats and 45 species of birds to England to be exhibited at meetings of the Zoological Club of the Linnean Society in 1828. Later William, sent a foetal specimen of a dolphin (Fletcher 1920).
While no papers dealing especially with Cuban insects were published by W.S. Macleay, among his papers were thirty nine water-colour drawings of lepidopterous larvae, from which he may have reared adults. Besides these there are a number of pencil or pen and ink sketches of Lepidoptera, scorpions, ticks and mites (Fletcher 1920).
The scientific world of today has been given an opportunity to know what was on the Island of Cuba in the years 1825 to 1836 due to the scientific
Proc. Linn. Soc. N.S.W., 131, 2010
D. CROSS AND E. JEFFERYS
endeavours of William Sharp Macleay in the form of over 7000 dry pinned labelled insects now placed together as the Cuban insect collection are housed in the insect collection in the Macleay Museum at the University of Sydney.
ACKNOWLEDGEMENTS
In July 2009 Dominic Cross was awarded the Macleay Miklouho-Maclay Fellowship at the Macleay Museum. At this time his supervisor of the Fellowship was Ms Elizabeth Jefferys, who was the Curator of natural History at the Macleay Museum at the University of Sydney. We thank the Macleay Museum for giving us the opportunity to complete this catalogue. We appreciate the fact that most of the identifications of the Cuban insects were organized by Dr Woody Horning a Curator at the Macleay Museum from 1982 to 1994. Dr Woody Horning identified much of the insects himself and organized other American taxonomists to identify material as well.
REFERENCES
Naumann I.D. and Steinbauer M.J. (2001). Egg parasitoids of Australian Coreidae (Hemiptera). Australian Journal of Entomology 40, 9-16.
Triplehorn, C.A. and Johnson NF (2005) “Borror and DeLong’s Introduction to the Study of Insects.’ (Thomson Learning, Southbank, Victoria, Australia.)
Fletcher, J.J. (1920). The Society’s heritage from the Macleays. Proceedings of the Linnean Society of New South Wales 45, 567-635.
Macleay, W.S. (1838). ‘Illustrations of the Annulosa of South Africa’. (Smith, Elder and Co., London).
Proc. Linn. Soc. N.S.W., 131, 2010
D9.
THE MACLEAY COLLECTION OF CUBAN BEETLES
CATALOGUE Blattodea FAMILY NUMBER Sitio A oe al Ae eciaaed cca =a | a Maa sei
tO Wn
oo
Coleoptera FAMILY
NUMBER 5 4 eae eee ee wha
iar er 0 r ea one chal ads Ba )
tO
4
— iw) — ee ps i)
8 2 2
Carabidae apie LT Es aa ETE Cerambycidae | Amphidesmus | | | Cerambycidae | Callichroma | | Cerambycidae Cerambycidae Eburia (orale nas thee | Cerambycidae | Eburodacrys |
—
—=}h
Cerambycidae | Elateropsis Cerambycidae Elateropsis fuliginosa Fabricius
Cerambycidae venusta Chevrolat
Cerambycidae Cerambycidae | Elateropsis |
ee Le ate Cerambycids [Leptosptis | FCerambyeidae | Odontacea [| |
Cerambycidae maculicornis Chevrolat 1862 Cerambycidae
30 Proc. Linn. Soc. N.S.W., 131, 2010
Cerambycidae Stenodontes Cerambycidae
Chrysomelidae Cassida Chrysomelidae Coptocycla Chrysomelidae
Ciidae
Curculionidae Attelabus
Curculionidae Baridius madrimaculatus Boheman
Curculionidae Calandra
Curculionidae
Curculionidae Eurhinus
Curculionidae Exophthalmus Curculionidae Exophthalmus Curculionidae Exophthalmus Curculionidae Exophthalmus
Curculionidae Exophthalmus
Curculionidae Hilipus Curculionidae Hilipus | Curculionidae Hilipus
Curculionidae Lachnopus Curculionidae Lachnopus
Curculionidae Lachnopus
D. CROSS AND E. JEFFERYS
damicornis Linnaeus 1771 5 161
dorsopunctata Boheman 16 2
27
6
4
2
agaves 1
sericea Olivier 1807
Curculionidae 14
i
i)
pEmonee jee sl haat ae alee Od] uous |
Jreyreissi Boheman 1836 guttatus Boheman 1843 rusticus Boheman 1836
curvipes Fabricius
N |W] tv
i
hispidus Gyllenhal
vittatus Gyllenhal
Curculionidae Lachnopus Curculionidae Pachnéus
Curculionidae Pachnéus
Curculionidae Peltophorus
Curculionidae Polydacrys
Sle IVNIniIwlyel|wsy
—
azurescens Gyllenhal
litus Germar
NO] nn] dy
modestus Gyllenhal
Curculionidae Prepodes Curculionidae Ptilopus
Curculionidae Rhina
Curculionidae Scyphophorus
Curculionidae Sphenophorus
Curculionidae Sphenophorus
Curculionidae Tetrabothynus
Curculionidae
Curculionidae Curculionidae Curculionidae
Dytiscidae Rhantus Dytiscidae
Tetrabothynus
Tylomus Xyleborus
Elateridae Pyrophorus
Histeridae
Lampyridae
Lycidae Calopteron ite | Mordetlidae |
3 9 Elateridae 2 1 5 ;
spectabilis Dejean 14
vittatus Dejean
9 scrutator Olivier 4 1
atheniunus Schedl
sericeus Latreille
spectabilis Gyllenhal
calidus Fabricius 1792
phosphorescens
bicolor Linnaeus i
lon
Prog, Eimns Soc, NESW. 131, 2010
31
THE MACLEAY COLLECTION OF CUBAN BEETLES
Passalidae Passalus Passalidae Passalus
Trox | Trogossitidae | UNIDENTIFED |
Diptera
ea ae iain I a [ONINENTREED|Ser PUNIDENTIFIED | |_|
Hemiptera
GENUS | SPECIES | NUMBER BBelasiomaticize) | ee aa | ae Cees ae OC ae a ae ee 2 Cicadidae Cicada viridicincta Macleay 6 KE ae a a a ee sexi ae ea a eee Resa «Eo ee aE Se Cascio s(t ee as Ee or) SGennitize;etien "5 Eeiiseg)|_ 1Ab nent Daa tiie Gon Caen Membracidae
cS ee Er ee ea Rec ae es
es aS a
Rio a
Gascon || cane ee eee Pentatomidae ee ee ee ee
Redividacle) )/|Uzioaa | es eS ee eee
| Reduviidae | UNIDENTIFIED
ened, ieee | ENDEMED |e eoe | eee | omen
52. Proc. Linn. Soc. N.S.W., 131, 2010
D. CROSS AND E. JEFFERYS
FAMILY Anthophoridae Apidae
Apidae
ethylide os
@)
2 S) S AS) g
ENUS SPECIES NUMBER irate cere we
jimbriata Fabricius 1804
a) < 3 @Q i=) lo} as} — Q 5 i)
Braconidae
Chalcididae Brachymeria Chalcididae Brachymeria Chalcididae Brachymeria robusta Cresson 44 Chaleididae | Brachymeria | 47 Chaleididae 8 Chalcididae 3 debilis Say 1836 4 xanticles (Walker) 8 ARIE (CESSED) 19 Chalcididae Spilochalcis femorata (Fabricius) 8 Chalcididae Spilochalcis maniae (Riley) 2 | Chalcididae Spilochalcis Chalcididae Spilochalcis
3 Chalcididae Spilochalcis 10 Chalcididae Spilochalcis
1 110 3 1
3 0
— —
Chalcididae Spilochalcis Chalcididae
Chrysididae Caenochrysis 1 Chrysididae Chrysis insularis Guérin 5 Chrysididae Chrysis 5 Chrysididae Chrysis 15
Chrysididae | Chrysis [| purpuriventris | | Chrysididae | Chrysis
Chrysididae Holopyga ventralis Say 2) Chrysididae 1 (pete) Cynipidae a a a TE
ee | aE [Huser | se ew eee 2 es a TST Geet a a ry
Formicidae
Formicidae 13
Formicidae Crematogaster 1 Formicidae Cyphomyrmex 1 Formicidae Odontomachus relictus 13
Camponotus
Proc. Linn. Soc. N.S.W., 131, 2010
THE MACLE
Megachilidae ii Mutillidae
Platygastridae
Pompilidae Pepsis
Pompilidae
Scoliidae
AY COLLECTION OF CUBAN BEETLES
trifasciata Burmeister
trifasciata Burmeister
Scoliidae Elis Sphecidae Monedula
Sphecidae Nysson
Sphecidae Nysson
insularis Dahl
albilabris
collaris
Nysson
Sphecidae
Nysson
hyalius
sericeus
Sphecidae
Sphecidae
Tiphiidae
Vespidae Ancistrocerus | Vespidae | Eumenes Vespidae Euodynerus
Pachodynerus [| | ST
9
cingulatus Cresson
7 2 50
Parancistrocerus | enyo (Lepeletier) 1841 27], 12
Vespidae Zeta pe Vespidae Zethus 14 Lepidoptera FAMILY GENUS SPECIES Lycaenidae Cyclargus ammon (Lucas) 1857 Lycaenidae Eumaeus atala Poey 1832 ] Lycaenidae Leptotes theonus (Lucas) 1857 Lycaenidae Lycaenidae Nymphalidae | Anaea Nymphalidae Apatura pavonii Latreille 3 Nymphalidae Eunica 2 Nymphalidae Hypanartia | paullus Fabricius 1793 2) Nymphalidae Megalura Nymphalidae Metamorpha IENpEERCRE[Eurcioddes | Nymphalidae | Siderone | Nymphalidae
Lo
4 Proc. Linn. Soc. N.S.W., 131, 2010
D. CROSS AND E. JEFFERYS
Papilio Papilio Papilionidae Papilio
Papilionidae Papilio
Papilionidae Papilio - ey Db OuUVd Pieridae
Sphingidae UNIDENTIFIED
UNIDENTIFIED
Neuroptera
FAMILY Myrmeleontidae
UNIDENTIFIED
_ Odonata
as 200
Orthoptera
iran imam) | NUMBER | [psa
Phasmatodea
FAMILY GENUS | SPECIES | NUMBER
UNDENDIED | ot. | 20
eee | Pulicidae | Ctenocephalides | felis (Bouché) 1836 | 8 | [Tee PE a ee Te
Proc. Linn. Soc. N.S.W., 131, 2010
oe LAP tts ORGANS Hi o11,.4 ee i aha | om * —_———— ipoiny, ‘ae r Diner @ ee | eer ‘ : psn My \ La vr eet, bh yea 5 — age Saal a gtr Gem b+ IE | | Np means ig ; SH, eS ee ee ie ati ¢ oy ; Chey Gaps) qn: 0 io at So
re oe *Y {ahi A LerwmT}* y [et & naan oy
|
e ‘ r ; _ ai i ® ' : ase | 4 nya + oye Nic mae ae Seen? Be 2 a —~
- 7 a * ul 7 - % a vi — - ee \ _ > _— ee } : at a! i ‘ie < ; t ; an ih, meer tr ale ay See ‘ Tey 4 is { er , ; TSM (oT ase) RU Tae ae 1 — rer © a ee i T F i Li i j 4 , hi gt —— - i nleermie f Laie | ti £0 ai Thay : ‘ ! i i x Py i i prey cP "pee on ay slat Car 5 ) i 1 = . APES) J i } i : a 4,10 a) ¥ J i i gi i < => > ia ’ oO ) r oo | es “ 7 ” - : , V bind 1 i i i ' . N i i ; ke if Ti nl r i) » ¢ m I \ jaa uy 7 Fr) A amy Bes 1 te a i Ve ' iis = Fu ees 5 fhe ie Blea ‘haw } " \ ry ‘ i Al a P 7 fi i a: i ] a fa t - ay oe bs) bok aed: ee ete : Tr fom : D a a Aan i ea cel ‘ P r “ ai pte a rey Pik ha ie i ey ' ae PORES J} Pe anne } : Pia —_ . } tr
— tg at
Description of a New Species of Inola Davies (Araneae: Pisauridae), the Male of I. subtilis Davies and Notes on Their Chromosomes
MartIN Tio! AND MARGARET HUMPHREY~*
'Faculty of Medicine, University of Sydney, NSW 2006, Australia *Australian Museum, 6 College Street, NSW 2010, Australia *Corresponding author (margaret.humphrey@yahoo.com.au)
Tio, M. and Humphrey, M. (2010). Description of a new species of Inola Davies (Araneae: Pisauridae), the male of Z. subtilis Davies and notes on their chromosomes. Proceedings of the Linnean Society of
New South Wales 131, 37-42
A new pisaurid spider, Inola daviesae sp.n. is described from northern Queensland together with the first description of the male of /. subtilis. The meiotic chromosomes of both species are discussed.
Manuscript received 5 March 2010, accepted for publication 21 April 2010.
KEYWORDS: chromosomes, /no/a, Pisaurid, Queensland, rainforest, spider.
INTRODUCTION
Australian pisaurid spiders are generally not web builders, except for members of J/no/a Davies, 1982 and Dendolycosa Koch, 1876. The genus Jnola includes three species from northeastern Queensland ( Davies, 1982). Like Davies’ Inola species, Inola daviesae sp.n. described here is a delicate, medium- sized spider associated with tropical rainforest. As with other members of the genus, this spider runs on the upper surface of its horizontal sheet web. These webs project from the trunks of rainforest trees or embankments. A short silken funnel extends from the sheet web to a retreat in a tree trunk or embankment. The females, like those of other pisaurids, grasp their egg sacs in their chelicerae when disturbed and carry them into their retreat (Davies, 1982).
Abbreviations: CL cephalothorax length; CW cephalothorax width; AL abdomen length; AW abdomen width; MOQ median ocular quadrangle; AM Australian Museum; QM Queensland Museum.
MATERIALS AND METHODS
Morphology
Measurements were made with an ocular micrometer and converted to millimetres. Measurements are for a single specimen with a range of variation if significant. Spines have been recorded as number per surface for each segment, as they were often staggered.
Chromosomes
Live penultimate male spiders were anaesthetized with CO,. The testes were dissected out and sections were spread, fixed and stained after the method of Rowell (1991). These preparations were viewed and photographed using a light microscope. Counts and other observations were noted from photographs of many (>50), suitable meiotic cells in metaphase and chromosome numbers for species determined by the mode.
NEW SPECIES OF JNOLA
SYSTEMATICS Genus /nola Davies, 1982
Inola Davies, 1982: 479 Type species: /nola amicabilis Davies, 1982, by original designation (page 480).
Inola daviesae n.sp. Figs (1-4, 7-11)
Types
Holotype: male, Leo Creek, Macllwraith Ranges, North Qld. [13°32’S 143°29’E], July, 1995, M. Humphrey, M. Moulds, KS58316 (AM). Paratypes: 1 female, same data as holotype, KS58322 (AM); 1 male, 5 females, Qld. MaclIlwraith Ranges, Leo Creek [13°32’S 143°29°E], 20 Jul 1995, M. Humphrey, M. Moulds, F. MacKillop, KS43933 (AM); 1 male, 1 female, data as for holotype, QMS 83903 (QM).
Other material examined Eleven juveniles, same data as holotype, KS58315 (AM).
Distribution Rainforest, MaclIlwraith Range, north-eastern Queensland at an altitude of approximately 500m.
Diagnosis
Males can be distinguished from other members of the genus by the distinctive spannerhead-shaped distal portion of the median apophysis of the male palp (Fig. 8). The female scape is narrow while that of I. cracentis is broad and that of /. subtilis is triangular, pointed posteriorly and broad anteriorly.
Description of male
Measurements of holotype: CL 4.3, CW 2.7, AL 5.9, AW 1.7. Eye group: anterior width 1.1; posterior width 1.1; length 0.6; MOQ: anterior width 0.4; posterior width 0.5; length 0.5. Maxilla: length 1.3; width 0.8; Sternum: length 1.9; width 1.9; Colulus: length 0.2; width 0.3. Leg lengths:
Palp i 2 3 4 Femur 4.5 11.9 11.4 7.6 eS Patella 1.6 1.9 2.0 1.4 eS Tibia 1.9 ital ES 8.9 10.3 Metatarsus
—_ Sal 14.6 9.8 15.4
Tarsus 4.6 4.9 4.6 3:5 4.9 Total 12.6 44.9 43.9 a2 43.4
38
Spine notation: Palp: femur, d3p1; patella, dSpI1r1; tibia, d2r2; tarsus, 0. Leg I: femur, d4p5; patella, d1; tibia, d3p3r2v3; metatarsus, p4r4v1, whorl of four small spines distally ; tarsus, 0. Leg II: femur, d2p5r5; patella, dl; tibia, d2p3r4v3; metatarsus, p3r4, whorl of four small spines distally; tarsus, 0. Leg III: femur, d2p4r5; patella, dl; tibia, p2r4v3; metatarsus, p4r4, whorl of four small spines distally; tarsus, 0. Leg IV: femur, d3p4r2; patella,d1; tibia p3r3v2; metatarsus, d4p4; tarsus,0. Note: four distal spines on end of each metatarsus.
Eye diameters roughly equal. Cephalothorax patterned (Fig.l). Abdomen with centrai pale stripe to almost half the length of abdomen. Pair of pale latero-dorsal stripes, running three quarters of the abdomen. Two or three pairs of prominent pale spots between the central and the latero-dorsal stripes. Legs banded.
Palp (Figs.7, 8). Digitiform portion half the length of the palpal tarsus. Median apophysis large and partly membraneous, partly sclerotised. Distal sclerotised portion bifid (spanner-like). Embolus slender and curved. Conductor behind median apophysis with a fold distally.
Description of female
Measurements of KS58322: CL 3.9, CW 3.4, AL 6.9, AW 4.7. Eye group: anterior width 1.5; posterior width 1.6; length 1.0; MOQ: anterior width 0.7; posterior width 0.8; length 0.7. Maxilla: length 1.6; width 1.0. Sternum: length 2.6; width 2.1; Colulus: length 0.2; width 0.3. Leg lengths:
Pal 1 2 3 4 Femur 2.8 8.6 9.4 Tes) 9.4
Patella 1.0 DD, 1.9 1.6 1.6 Tibia 1.4 8.8 93 6.1 8.0 Metatarsus
— 10.4 8.6 7.9 12.5 Tarsus 3.1 3.0 BS 3.1 43 Total 8.3 33.0 B2a/ 26.2 35.8
Spine notation: Palp: femur, d1p1; patella, d1p1; tibia d2p2rl, tarsus, p2. Leg I: femur, d2r2; patella, dl; tibia, dlr2vl; metatarsus, d3r4v2, whorl of four small spines distally, tarsus, 0. Leg II: femur, d2p5r5; patella, dl; tibia, dlp2r2vl; metatarsus, d3p2r4v2, whorl of four small spines distally; tarsus, 0. Leg III: femur, d4p2v1; patella, dilrl; tibia, 0; metatarsus, dlp2r2v2, whorl of four small spines distally; tarsus, 0. Leg IV: femur, d4r5; patella dirl; tibia, 0; Metatarsus, d2p3rlv2, whorl of four small spines distally; tarsus, 0. Note: four distal spines on end of metatarsus (every leg).
Proc. Linn. Soc. N.S.W., 131, 2010
M. TIO AND M. HUMPHREY
Figures 1-7. 1, Inola daviesae sp. n. male carapace, dorsal, (holotype). 2, Imola daviesae sp.n. male cepha- lothorax, lateral, (holotype). 3, Inola daviesae sp. n. epigynum, external, (KS58322). 4, Inola daviesae sp. n. epigynum, internal, ventral. 5, Inola subtilis, male palp, ventral, (KS58321). 6, Inola subtilis, expanded male palp, retrolateral, (KS58320).
Proc. Linn. Soc. N.S.W., 131, 2010 39
NEW SPECIES OF INOLA
30 jim 11 va 2opm 12
Figures 7-12. 7, male palp of Jnola daviesae sp.n. 8, median apophysis (ma), embolus (e) and conductor (c) of Inola daviesae sp. n. 9, Inola daviesae sp.n. female on sheet web. 10, Inola daviesae sp.n., prophase male meiotic chromosomes showing two dense sex chromosomes (arrowed). 11, Inola daviesae sp.n., male meiotic cell showing 14 pairs of chromosomes. 12, Inola subtilis, male prophase meiosis showing two densely stained sex chromosomes (arrowed).
40 Proc. Linn. Soc. N.S.W., 131, 2010
M. TIO AND M. HUMPHREY
Epigynum (Figs 3, 4). Scape a narrow bar. Insemination ducts arise near hind edge of the epigastrum and travel forward. Large stalked spermathacae. Insemination duct enters near the base of the posterior spermathacae (fertilisation duct leaves below this junction).
Chromosomes
For males of /. daviesae sp. n., 2N = 28 (Fig. 11), including two subequal, darkly staining sex chromosomes. Most of the 13 pairs of autosomes in Inola daviesae sp. n. appear to be telocentric. The two sex chromosomes are easily distinguished in prophase of meiosis (Fig. 10). They migrate from the equator of the spindle in metaphase as a pair and earlier than the autosomes. Such sex chromosomes and their behaviour have been observed in other spiders by Rowell (1991). According to a survey of spider chromosome studies, (Rowell, personal comm.), female spiders have double the number of _ sex chromosomes to those of the male. Presuming this species follows the same sex determination mechanism, males of /nola daviesae n. sp. would be XX and females XXXX, giving females 2N = 32.
Etymology. Named for Valerie Todd Davies who described the genus.
Inola subtilis Davies, 1982 (Figs 5, 6)
Material examined
1 male, Goldsborough S. F., Qld., July, 1995, M. Humphrey, KS58321 (AM); 1 male, data as for KS58321, QMS83902 (QM); 3 males, data as for KS58321, KS58320 (AM); 1 male, Palm Cove, FNQ, J.Olive, 6 Sept 1995, sheet web on fallen log, KS044108 (AM); Goldsborough Valley SF, rainforest strangler fig, 27 Jul 1995, M. Humphrey, KS043900 (AM).
Distribution
Material from Davies’ description of the species indicates a distribution on the western edge of suburban Cairns. The material examined above extends this distribution from Palm Cove (north of Cairns) to the Goldsborough Valley in the south.
Diagnosis for male
Unlike the other three members of the genus, the sclerotised distal portion of the male palpal median apophysis forms two, fused, parallel, curved processes
Bro. mn, Soc. N:SAW.A3 15 20110
(Fig. 5). Proximally is a long, narrow sclerotised spur pointing ventrally, at right angles to the palp. Conductor sclerotised, retrolateral, behind the large median apophysis and bearing a spine distally.
Description of male
Measurements of KS58321: CL 3.5, CW 2.8, AL 4.4, AW 1.44; Eye group: anterior width 0.8; posterior width 1.2; length 0.8; MOQ: anterior width 0.5, posterior width 0.6, length 0.5. Maxilla: length 1.0; width 0.5. Sternum: length 1.8, width 1.7. Colulus: length 0.3, width 0.5. Leg lengths:
Palp 1 2 3 4 Femur 2.0 10.0 9.3) es) 9.4
Patella 0.6 1.6 1.6 1.5 1.5 Tibia 0.8 10.3 9.4 6.9 8.9 Metatarsus
— 12.6 12.1 9.1 13.0 Tarsus 1.6 3.9 3.6 2.5 353) Total 5.0 38.4 36.0 DS 36.1
Spine notations: Palp: femur, d2p1; patella, dipIirl; tibia, d2p2; tarsus, 0. Leg I: femur, d2p8r3; patella, dl; tibia, d3p2r2v4; metatarsus, d3p2r5v2; tarsus, 0. Leg I: femur, p5; patella, d5p6r3; tibia, d2p2r2v3; metatarsus, dlp3r3v1; tarsus, 0. Leg III: femur, d2p5r5; patella, d1; tibia, d2p3r3v3; metatarsus, d2p2r2v2; tarsus, 0. Leg IV: femur, d2p5r2; patella, d2p4r2v3; tibia, dlplr3vl; metatarsus, d2p2r2; tarsus, 0.
Abdomen long and narrow. Abdominal pattern with pale centre stripe and a pair of pale latero-dorsal stripes. Pairs of prominent pale spots as in 1. daviesae but spots continue in line and merge to form a pair of additional stripes. Legs banded.
Male palp (see diagnosis): Length of digitiform portion almost half of palpal tarsus. Embolus curved, slender, lying between median apophysis and conductor.
Chromosomes
Because of poor spreading, the number of chromosomes of J. subtilis could only be estimated. However, it is between 26 and 32 and most of the chromosomes are telocentric. There are two sex chromosomes (Fig. 12) and like those of 1. daviesae sp. n., they are darkly staining and migrate from the equator of the spindle earlier than the autosomes.
41
NEW SPECIES OF INOLA
ACKNOWLEDGMENTS
We are grateful to the following staff and departments of the University of Sydney; Assoc. Prof. L.W. Burgess, Dean, Faculty of Agriculture, for provision of a mentorship to the senior author; Assoc. Prof. H. A. Rose, Department of Crop Sciences and the Electron Microscope Unit for the use of their facilities. Our thanks also to Dr Valerie Todd Davies, Queensland Museum, for specimen identifications and to Dr M. Gray and Dr M.S. Moulds, Australian Museum, for
valuable advice and assistance. REFERENCES
Davies, V. T. (1982). /nola nov. gen., a web-building pisaurid (Araneae: Pisauridae) from northern Australia with descriptions of three species. Memoirs of the Queensland Museum 20(3): 479-487.
Rowell, D.M. (1991). Chromosomal fusion in De/ena cancerides (Araneae: Sparassidae). I. Chromosome pairing and X-chromosome segregation. Genome 34: 561-573.
42 Proc. Linn. Soc. N.S.W., 131, 2010
A Late Ordovician Conodont Fauna from the Lower Limestone Member of the Benjamin Limestone in Central Tasmania, and Revision of Tasmanognathus careyi Burrett, 1979
Y.Y. ZHEN!, C.F. BurretT’, I.G. PERCIVAL? AND B.Y. Lin*
‘Australian Museum, 6 College Street, Sydney, N.S.W. 2010, Australia (yongyi.zhen@austmus.gov.au); *School of Earth Sciences, University of Tasmania, GPO Box 79, Hobart, Tasmania 7001, Australia (cliveburrett@gmail.com);
*Geological Survey of New South Wales, 947-953 Londonderry Road, Londonderry, N.S.W. 2753, Australia (ian.percival@industry.nsw.gov.au);
“Institute of Geology, Chinese Academy of Geological Sciences, Beijing, China, 100037.
Zhen, Y.Y., Burrett, C.F., Percival, I.G. and Lin, B.Y. (2010). A Late Ordovician conodont fauna from the Lower Limestone Member of the Benjamin Limestone in central Tasmania, and revision of Tasmanognathus careyi Burrett, 1979. Proceedings of the Linnean Society of New South Wales 131,
43-72.
Ten conodont species, including Aphelognathus? sp., Belodina compressa, Chirognathus tricostatus
sp. nov., Drepanodus sp., gen. et sp. indet., Panderodus gracilis, Protopanderodus? nogamii, Phragmodus undatus, Tasmanognathus careyi and T. sp. cf. T. careyi are documented from the Lower Limestone Member of the Benjamin Limestone, Gordon Group, exposed in the Florentine Valley and Everlasting Hills region of central Tasmania. For the first time since its establishment three decades ago, the type species of Tasmanognathus, T. careyi, is revised with recognition of a septimembrate apparatus including makellate M, alate Sa, digyrate Sb, bipennate Sc, tertiopedate Sd, carminate Pa, and Pb (angulate Pb! and pastinate Pb2) elements. Co-occurrence of Phragmodus undatus and Belodina compressa in the fauna indicates a latest Sandbian to earliest Katian (Phragmodus undatus conodont Zone) age for the Lower Limestone Member of the Benjamin Limestone. All species previously attributed to Zasmanognathus are briefly reviewed, and the distribution of the genus is shown to be more widespread than hitherto recognised (in New South Wales, North China, Tarim Basin, South Korea and northeast Russia), with a probable occurrence in North
American Midcontinental faunas.
Manuscript received 16 September 2009, accepted for publication 26 May 2010.
KEYWORDS: Benjamin Limestone, biogeography, biostratigraphy, conodonts, Late Ordovician,
Tasmania, Zasmanognathus.
INTRODUCTION
Ordovician conodont faunas of Tasmania are relatively poorly known in comparison to those from the mainland of Eastern Australia. Only three papers — Burrett (1979), Burrett et al. (1983) and Cantrill and Burrett (2004) — have dealt systematically with a small number of species. The present contribution, which describes the comparatively diverse fauna from the lower part of the Benjamin Limestone, is the first part of a revision of all known conodonts from
Tasmania. This project aims to provide a firm basis for conodont-based correlations of the carbonate- dominated Gordon Group with limestones along the Delamerian continental margin in New South Wales, with strata in offshore island arc settings in central N.S.W. (Macquarie Arc), and with isolated limestone pods in the New England Orogen in northeastern N.S.W. and central Queensland.
Given the rarity of graptolites in the predominantly shallow-water platformal succession forming the Delamerian margin succession, and the sparsely documented occurrences of conodonts,
LATE ORDOVICIAN CONODONTS FROM TASMANIA
biostratigraphical zonation in Ordovician rocks of Tasmania is currently largely reliant on shelly macrofossils. Banks and Burrett (1980) established a series of twenty successive faunas (designated OT assemblages 1-20), one of which (OT 12) was defined by the occurrence of several conodont species including Tasmanognathus Chirognathus monodactyla, Erismodus gracilis and Plectodina aculeata in the basal Benjamin Limestone. This fauna (based at the time on unpublished studies by Burrett, with no species illustrated or described in the 1980 paper) is revised here. Our study has not identified the last two named species, and has recognised a new species of Chirognathus in place of C. monodactyla. Burrett (in Webby et al. 1981, p.12) summarised the occurrences of conodonts in the Tasmanian Ordovician succession. He noted the first appearance of the biostratigraphically important species Phragmodus undatus in strata immediately above the Lords Siltstone Member in the middle of the Benjamin Limestone; however, our reassessment of the fauna has identified the presence of this species in the underlying lower part of the Benjamin Limestone. Laurie (1991) defined an alternate series of 20 faunal assemblages based on Tasmanian brachiopods, ranging in age from Early Ordovician (Tremadocian) to earliest Silurian. Where possible, these brachiopod faunas were tied in to conodont occurrences, mainly derived from Burrett’s (1978) unpublished thesis studies.
A biogeographically significant component of the Tasmanian conodont fauna is 7asmanognathus Burrett, 1979, which was first identified from the Lower Limestone Member of the Benjamin Limestone exposed in the Florentine Valley and Everlasting Hills region of central Tasmania (Fig. 1). This genus has subsequently been widely recognized as occurring in rocks of early Late Ordovician (Sandbian) age in eastern Australia and China. Low yields (averaging two specimens per kg) of conodonts from the Gordon Group carbonates collected and processed by Burrett (1978) resulted in Jasmanognathus being imperfectly defined. Thirty years after its initial documentation, revision of the type species, 7. careyi Burrett, 1979 has become urgently needed in order to better understand its multielement apparatus, phylogenetic relationship and precise stratigraphic range in the type area. The purpose of this paper is to describe the conodont fauna from the middle part of the Gordon Group in the Settlement Road section of the Florentine Valley area, equivalent to the level yielding Tasmanognathus, based on five recently collected bulk samples of limestone totalling 49.5 kg that on dissolution in acetic acid have yielded an average
careyl,
44
of six elements per kg. These additional collections are supplemented by re-examination of Burrett’s original material including types and topotypes of T. careyi, and for the first time all the accompanying conodont fauna is documented by description and/or illustration, including Aphelognathus? sp., Belodina compressa (Branson and Mehl, 1933), Chirognathus tricostatus sp. nov., Drepanodus sp., Panderodus gracilis (Branson and Mehl, 1933), Protopanderodus? nogamii (Lee, 1975), Phragmodus undatus Branson and Mehl, 1933, and gen. et sp. indet.
REGIONAL GEOLOGIC AND BIOSTRATIGRAPHIC SETTING
Platform sedimentary rocks of the Early Palaeozoic Wurawina Supergroup, that are widespread in the western half of Tasmania, consist of the Late Cambrian — Early Ordovician Denison Group (mainly siliciclastics), conformably overlain by the Gordon Group (predominantly carbonates of Early to Late Ordovician age), in turn conformably or disconformably overlain by the Hirnantian (latest Ordovician) to mid-Devonian Eldon Group, which consists mainly of siliciclastics (Burrett et al. 1984; Laurie 1991). The Gordon Group attains a thickness of 2100m of carbonates and minor siltstones in its redefined type section in the Florentine Valley where it is divided into three limestone formations. The uppermost of these, the Benjamin Limestone, is divided into two limestone members (Upper and Lower) separated by a thin but regionally extensive, macrofossiliferous siltstone member (Lords Siltstone Member). The Benjamin Limestone predominantly consists of interbedded microcrystalline peritidal dolomitic micrite, dolostone and calcarenite with a maximum thickness of about 1200m. Some 400 conodont samples were initially collected over a Sm interval by Burrett (1978) from the various localities of the Gordon Group, but many of these samples were barren or had a very low yield, due to the peritidal to shallow subtidal depositional setting and high rate of sedimentation in the tropical shelf environments. Continuous efforts in the last 30 years by post- graduate students and academic staff of the University of Tasmania have accumulated significant amounts of conodont material for the age determination and biostratigraphic analysis of the Gordon Group (Burrett 1979; Burrett et al. 1983, 1984; Cantrill and Burrett 2004).
Carbonates that are coeval with the Lower Limestone Member of the Benjamin Limestone occur in many sections in northern, western and southern
Proc. Linn. Soc. N.S.W., 131, 2010
Y.Y. ZHEN, C.F. BURRETT, I.G. PERCIVAL AND B.Y. LIN
eEverlasting Hills Florentine e
Valley § p np aa
* Sample locations EG Gordon Group ee
Lanes 4Peak 0
Gells 4 Lookout
MT FIELD
a Mt Field West %
ides %Z. nun ee
AS?” NATIONAL % PARK
Florentine
i: > an
River \ NS loidg ~~ rt
Sandstone Sandstones (Early? Ordovician) Mabon (late Ordovician to Silurian?)
Figure 1. Maps showing the studied areas in central Tasmania and sample locations. A, Map of Tasma- nia showing the locations of Florentine Valley and Everlasting Hills (from Burrett 1978, 1979); B, Map showing the Florentine Valley area and sample location of the Nine Road Section (modified from Laurie 1991); C, Map showing the Settlement Road Section of the Florentine Valley and sample locations (modi- fied from Laurie 1991); D, Map showing Everlasting Hills area and sample location (from Burrett 1978).
Tasmania, but the Zasmanognathus careyi fauna has only been definitely found in the Florentine Valley and in the Everlasting Hills. The Florentine Valley sections (Figs | and 2) are found in the eastern side of a mid-Devonian synclinorial structure. This area was first mapped geologically by Corbett and Banks (1974) and because of its completeness, has subsequently been the focus of numerous palaeontological and sedimentological studies. However, active timber logging in this area has meant that some sections are now inaccessible, having been replanted with dense, almost impenetrable, forest.
The Everlasting Hills section (Fig.1D) was discovered in remote and moderately dense to thick
Proc. Linn. Soc. N.S.W., 131, 2010
vegetation and mapped by Ian McKendrick and Clive Burrett in 1975 (Fig.1D). This doline and cave-rich area has since been included in the South West Tasmania World Heritage wilderness area, and has undergone extensive regrowth so that it is now extremely difficult to access. The palaeotropical limestones in the Everlasting Hills are identical to those in the Lower Limestone Member of the Benjamin Limestone in the Florentine Valley, and consist of 3-6m thick Punctuated Aggradational Cycles (Goodwin and Anderson 1985) of mainly dolomitised, intertidal micrites with tidal channels and top beds containing a lower intertidal to high subtidal macrofauna. Somewhat deeper water, coeval carbonates (the Ugbrook Formation) occur in
45
ORDOVICIAN CONODONTS FROM TASMANIA
» 4
LATE
FLORENTINE VALLEY SETTLEMENT ROAD
FLORENTINE VALLEY
EVERLASTING HILLS
NINE ROAD
tAaieo 1 39 ‘ds snyjeuBouewse; en & iAaieo snyjeuBouewse) a &
snjepun snpowBeiyd ee timeBou gsnposapuedojosg eeep siioei6 snposepueg eS &
yepui “ds ja -uag @ ‘ds snpouedaiq ; e< snjejsoou) snyjeuBouiyD ORR essaidwoo eulpojag ry
‘ds gsnyjeubojoydy e
LORDS SILTSTONE MEMBER CASHIONS CREEK LIMESTONE
tAaueo jy (yo ‘ds snyjeuBouewse; e iAeseo snyjeuBouewsey ee snjepun snpowBeiy4 e—e sijoeiB snposepueg eo—_e ‘ds snpouedaig e snjejsoouy snyjeuBosy9 ee essaidwos eulpojag e
thaieo | jo “ds snyjeuBouewse; e_eo0 ifaieo snyjeuBouewses e- ee snjepun snpowBbeiyd & ‘ds snpouedaiq ee = snjejsoou} snyjeuBouiyD e S ‘ds gsnyjeubojaydy @
uw a6 9 ¥ rn YaaWAW YaddN = one Zo ra a Baad ANOLSAWI1 SYaaWyvy Be iz YagW3W YaMO1 OSw 20 g°= so JNOLS3WI1 NINVPN3a = 5°
dnouS NOGYOS
Figure 2. Three stratigraphic sections showing the sample horizons and ranges of the conodont species in the Lower Limestone Member of the Benjamin Limestone, Gordon Group, in central Tasmania.
Proc. Linn. Soc. N.S.W., 131, 2010
46
YY ZHEN, CE BURREDIE EGSPERCIVAL AND BLY: LIN
northern and western Tasmania (Burrett et al. 1989) but these lack Tasmanognathus. This suggests that Tasmanognathus was mainly restricted to peritidal tropical environments in the Late Ordovician.
The TYasmanognathus fauna is associated with a strongly endemic macrofauna in the lower and middle parts of the Lower Limestone Member, Benjamin Limestone, including the brachiopods Lepidomena Laurie, 1991, Yasmanorthis Laurie, 1991 and the nautiloids Gorbyoceras settlementense Stait and Flower, 1985, Paramadiganella Stait, 1984 and Tasmanoceras zeehanense YVeichert and Glenister, 1952 (Laurie 1991; Stait 1988). Tasmanognathus careyi is found in two of the twenty Ordovician brachiopod assemblages (or biozones) recognised by Laurie (1991); the Zasmanorthis calveri and the younger Zasmanorthis costata assemblages.
AGE AND CORRELATION OF THE FAUNA
In the conodont fauna associated with Tasmanognathus careyi from the Lower Limestone Member of the Benjamin Limestone in central Tasmania, occurrence of Phragmodus undatus and Belodina compressa is crucial for age determination and regional correlation, as both species are cosmopolitan and age diagnostic. The former had a relatively long stratigraphic range, extending from the base of the Ph. undatus Zone (in the upper Sandbian) to the top of the Katian, and the latter first occurs at the base of the B. compressa Zone and extends to the base of the B. confluens Zone (Sweet 1988). Co-occurrence of these two species and absence of any diagnostic species of either the B. confluens or P. tenuis zones indicates a latest Sandbian to earliest Katian age (Phragmodus undatus Zone) for this Tasmanian fauna.
Chirognathus is also morphologically distinctive with the two previously-reported species (Chirognathus duodactylus Branson and Mehl, 1933 and Chirognathus cliefdenensis Zhen and Webby, 1995) restricted to the upper Sandbian- Katian interval (Sweet 1982; Zhen & Webby 1995). The new species from Tasmania described herein is morphologically similar to the type species of the genus, C. duodactylus Branson and Mehl, 1933. This species with a well-known multi-element apparatus is widely distributed in Sandbian strata of the North American Mid-continent ranging from the Pygodus anserinus Zone to the Phragmodus undatus Zone (Sweet in Ziegler 1991). The second species, Chirognathus cliefdenensis Zhen and Webby, 1995, occurs in a stratigraphically slightly younger interval
Proc. Linn. Soc. N.S.W., 131, 2010
in central New South Wales, where it is recorded from the upper Fossil Hill Limestone to the lower Vandon Limestone (early Katian) of the Cliefden Caves Limestone Subgroup (Zhen and Webby 1995), from the Downderry Limestone Member (late Katian) of the Ballingoole Limestone of the Bowan Park Limestone Subgroup (Zhen et al. 1999), and from allochthonous limestones of Katian age emplaced in the Silurian Barnby Hills Shale (Zhen et al. 2003a). The Lower Limestone Member of the Benjamin Limestone exposed in the Everlasting Hills and Florentine Valley areas in central Tasmania is the type stratum of Yasmanognathus careyi Burrett, 1979. Since the initial documentation of this species, at least ten additional species from lower Sandbian to upper Katian strata predominantly of North China and eastern Australia have been accommodated in Tasmanognathus (see Systematic section for further discussion). The origin and phylogenetic relationships of Yasmanognathus remain uncertain as most of these species were poorly documented and need to be revised. Reassessment of 7. careyi herein suggests that Zasmanognathus may be closely related to so- called “Ordovician ozarkodinides” (Sweet 1988, p. 91-92), an informal group including forms like “Plectodina’, Aphelognathus and Yaoxianognathus. Based on similarities of their general morphology and apparatus construction, Zasmanognathus, as a sister group, seems closely related to Yaoxianognathus. Tasmanognathus is potentially the direct ancestor of the latter, which was mainly restricted to eastern Gondwana and peri-Gondwanan terranes during the Late Ordovician (Katian). Strong biogeographic similarities (including Zasmanognathus) between the North China Terrane (or block) and eastern Australia were part of the evidence used by Burrett et al. (1990) to suggest that these blocks were contiguous or closely proximal during the Ordovician. Tasmanognathus was widely reported from the Sandbian in North China with recognition of three biozones based on the inferred lineage of Tasmanognathus species (An and Zheng 1990; Lin and Qiu 1990), from the oldest 7: sishuiensis Zhang in An et al., 1983 from the upper Fengfeng Formation (lower Sandbian), to 7. shichuanheensis An in An et al., 1985 from the middle-lower part of the Yaoxian Formation (upper Sandbian), and then to the youngest 7: multidentatus An in An and Zheng, 1990 (the latter is a nomem nudum, equivalent to T. borealis An in An et al. 1985; see Systematic Section for further discussion) from the upper part of the Yaoxian Formation (upper Sandbian-lower Katian). An and Zheng (1990, p. 95, text-fig. 9) illustrated the morphological changes from T. sishuiensis with a
AT
LATE ORDOVICIAN CONODONTS FROM TASMANIA
robust cusp and small, widely spaced denticles on the processes of the S elements, to 7) mu/tidentatus with a small, indistinct cusp in the Pa element and closer
the S elements. Importantly, similar morphological changes have also been observed between the two species of Zasmanognathus recognized in the Lower Limestone Member of the Benjamin Limestone in central Tasmania. A species described herein as 7. sp. cf. 7. careyi that bears a prominent cusp in the Pa element and small, widely spaced denticles on the processes of the S and Pb elements is more comparable with 7: shichuanheensis from the middle- lower part of the Yaoxian Formation, whereas 7: careyi with a small or indistinct cusp in the Pa element and long, closely spaced denticles on the processes of the S elements is closer to 7: multidentatus from the upper part of the Yaoxian Formation. 7. careyi was also reported from the middle part of the Yaoxian Formation in association with 7. shichuanheensis and Belodina compressa in Bed 3, about 44 m below the first occurrence of 7. multidentatus (An and Zheng 1990, p. 86-87), although An’s identification cannot be confirmed without re-examination of the original material (An et al. 1985) and further investigations. Occurrence of Taoqupognathus blandus at the top of the Yaoxian Formation in the Taoqupo Section of Yaoxian County (formerly Yaoxian; An and Zheng 1990) suggests that the Yaoxian Formation
may well extend to the lower Katian. Therefore, the morphological characters shown by the two species of Tasmanognathus from the Lower Limestone Member of the Benjamin Limestone support a correlation between this limestone unit in central Tasmania, and the middle part of the Yaoxian Formation in North China (with the possible occurrence of T. careyi), which An and Zheng (1990, p. 92, table 2) correlated with the C. wilsoni graptolite Zone (late Sandbian).
An and Zheng (1990, p.115) suggested that the Llandoverian conodonts illustrated by Lee (1982) from the Hoedongri Formation in the Taebaeksan Basin, Kangweon-Do of South Korea were comparable with the Tasmanognathus sishuiensis assemblage from the upper Fengfeng Formation of North China. In fact, in their revision of Lee’s original identifications (An and Zheng 1990, table 5, pp. 118- 119), they believed what Lee (1982) illustrated as Pterospathodus celloni (Walliser) should belong to Tasmanognathus sishuiensis, and considered that the Hoedongri Formation should be correlated with the Baduo Formation or the upper part of the Fengfeng Formation (Sandbian) of North China.
MATERIAL AND SAMPLING LOCALITIES
The current study is based on 683 identifiable specimens from 10 samples (See Table 1). Of these,
Table 1. Distribution of conodont species in the samples studied.
N
2 = species Aphelognathus? sp. Belodina compressa 8) Chirognathus tricostatus sp. nov. 36 6 Drepanodus sp. 20 Gen. et sp. indet. Panderodus gracilis All vep2 Protopanderodus? nogamii Phragmodus undatus Pas Seay) Tasmanognathus careyi 156 Tasmanognathus sp. cf. T. careyi 3 Total 290 15 48
Sera ate Shak SOs sy OF he ha 2 2 4
1 10
12 Digi leet ls oad 71 Pare ee sl 2, D5 3 3
4 3 2, 2 4 44 3 18 69
6 1 Sib Sil 116
209 265 26 Wian23 O5y— lilee2977 is} 4 if 14 2 34
2" 48" 26" 23 79936 120) 2) 683
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378 specimens are Burrett’s (1979) original material including types of Zasmanognathus careyi recovered from five samples collected from the Florentine Valley and Everlasting Hills sections (see Burrett 1979, p. 32, fig. 1 for sample locations and their stratigraphic horizons within the Lower Member of the Benjamin Limestone). Samples LLMB, C137 and C98 were collected from the Lower Limestone Member of the Benjamin Limestone exposed along the Nine Road (Fig. 1B). The Lower Limestone Member of the Benjamin Limestone is exposed as a 50m thick section (at Grid Ref. DP202157; 42°16.4’S, 146°2.65’E) to the north side of the Everlasting Hills (Fig. 1D). Two samples (JRC 2 and JRB) from this location produced relatively abundant conodonts (Table 1). The remaining 305 specimens were recovered from five large spot samples — YYF 1 (13 kg), YYF2 (8 kg), YYF3 (10 kg), YYF4 (7.5 kg), and YYFS5 (11 kg) — collected from the lower part of the Lower Limestone Member of the Benjamin Limestone in the Settlement Road section of the Florentine Valley area (Figs 1C, Ds
SYSTEMATIC PALAEONTOLOGY
All photographic illustrations shown in Figures 3 to 17 are SEM photomicrographs of conodonts captured digitally (numbers with the prefix IY are the file names of the digital images). Figured specimens bearing the prefix AM F. are deposited in the type collections of the Palaeontology Section at the Australian Museum in Sydney. All the syntypes except one (UTG96863 not located; figured by Burrett 1979, pl. 1, figs 17-18) and most of the other specimens of Zasmanognathus careyi illustrated by Burrett (1979) were relocated and made available for the current study. They have been now transferred to the Australian Museum collection, and a new AM F. registration number has been allocated to each of the specimens illustrated in this contribution.
The following species are documented herein only by illustration as they are either rare in the collection or have been adequately described elsewhere in the literature: Aphelognathus? sp. (Fig. 3J-K), Drepanodus sp. (Fig. 3C-F), gen. et sp. indet. (Fig. 3G-I), and Panderodus gracilis (Branson and Mehl, 1933) (Fig. 6A-I). Authorship of the new species Chirognathus tricostatus is attributable solely to Zhen. Taxa documented herein are alphabetically listed according to their generic assignment, with family level and higher classification omitted.
Proc. Linn. Soc. N.S.W., 131, 2010
Phylum Chordata Balfour, 1880 Class Conodonta Pander, 1856
Genus BELODINA Ethington, 1959
Type species Belodus compressus Branson and Mehl, 1933.
Belodina compressa (Branson and Mehl, 1933) Fig. 3A-B
Synonymy
Belodus compressus Branson and Mehl, 1933, p. 114, pl. 9, figs 15, 16.
Belodus grandis Stauffer, 1935, p. 603-604, pl. 72, figs 46, 47, 49, 53, 54, 57.
Belodus wykoffensis Stauffer, 1935, p. 604, pl. 72, figs 51, 52, 55, 58, 59.
Oistodus fornicalus Stauffer, 1935, p. 610, pl. 75, figs 3-6.
Belodina dispansa (Glenister); Schopf, 1966, p. 43, Ok I, ie, 7,
Belodina compressa (Branson and Mehl); Bergstr6m and Sweet, 1966, p. 321-315, pl. 31, figs 12-19; Sweet in Ziegler, 1981, p. 65-69, Belodina - plate 2, figs 1-4; Leslie, 1997, p. 921-926, figs 2.1-2.20, 3.1-3.4 (cum syn.); Zhen et al., 2004, p. 148, fig. 5A-I (cum syn.); Percival et al., 2006, fig. 3A-D.
Belodina confluens Sweet; Percival et al., 1999, p. 13, Fig. 8.21.
Material Ten specimens from two samples (see Table 1).
Discussion
Only compressiform (Fig. 3A) and grandiform (Fig. 3B) elements were recovered from the Tasmanian samples. These elements are identical with those recorded from the upper part of the Wahringa Limestone Member of the Fairbridge Volcanics (assemblage C, see Zhen et al. 2004), and others from drillcore samples in the Marsden district (Percival et al. 2006) of central New South Wales. Morphological distinction between B. compressa and closely related species, particularly B. confluens, was discussed by Zhen et al. (2004).
Genus CHIROGNATHUS Branson and Mehl, 1933
Type species Chirognathus duodactylus Branson and Mehl, 1D)3;3}-
49
LATE ORDOVICIAN CONODONTS FROM TASMANIA
Figure 3. A-B, Belodina compressa (Branson and Mehl, 1933). A, compressiform element, AM F.136480, JRC 2, inner-lateral view (1Y139-001); B, grandiform element, AM F.136481, JRC 2, outer-lateral view (TY 139-003). C-F, Drepanodus sp. C, Sb element, AM F.136482, JRC 2, outer-lateral view (1Y 139-005). D, Sb element, AM F.136483, JRC 2, inner-lateral view (1Y139-006). E, F, M element, AM F.136484, JRC 2, E, inner-lateral view ([Y139-004); F, basal view ([Y139-014). G-I, Gen. et sp. indet., all from YYF4, G, Sc element, AM F.136485, inner-lateral view ([Y136-022); H, ?P element, AM F.136486, outer-lateral view (IY 136-021); I, Sb element, AM F.136487, outer-lateral view (1Y136-019). J-K, Aphelognathus? sp. from YYF4, J, Pb element, AM F.136488, inner-lateral view (LY 135-025). K, Pa element, AM F.136489, inner-lateral view (1LY136-024). Scale bars 100 um.
revised the type species as having a seximembrate or septimembrate apparatus, and concluded that the 29 out of the 42 species recognized by Branson and Mehl (1933), Stauffer (1935), and others since the establishment of the genus could be confidently
Discussion
Chirognathus was established on 23 form species recognized by Branson and Mehl (1933, pp. 28-34, pl. 2) from the Harding Sandstone in Canyon City, Colorado with Chirognathus duodactylus as the : type species. Later Stauffer (1935) erected 15 form assigned to the genus, and in fact might belong to a species of Chirognathus from the upper Glenwood _ Single species apparatus of his revised C. duodactylus. Beds in the upper Mississippi Valley. Sweet (1982) He regarded 15 of Branson and Mehl’s (1933) and 13
50 Proc. Linn. Soc. N.S.W., 131, 2010
Y.Y. ZHEN, C.F. BURRETT, I.G. PERCIVAL AND B.Y. LIN
of Stauffer’s (1935) form species as junior synonyms of C. duodactylus, with the M element represented by form species C. duodactylus (= C. gradatus Branson and Mehl, 1933, = C. planus Branson and Mehl, 1933), Sa by form species C. multidens Branson and Mehl, 1933, Sb by form species C. panneus Branson and Mehl, 1933 (= C. isodactylus Branson and Mehl, 1933), Sc by form species C. eucharis Stauffer, 1935, Pa by form species C. varians Branson and Mehl, 1933 (= C. alternatus Branson and Mehl, 1933), and Pb by form species C. monodactylus Branson and Mehl, 1933 (= C. reversus Branson and Mehl, 1933). As defined by Sweet (1982, p. 1039), C. duodactylus has a ramiform-ramiform species apparatus including a bipennate M element with a short and laterally deflected anterior process and a long posterior process, an alate Sa element with a straight, laterally extended lateral process on each side, a digyrate Sb element varying from subsymmetrical (with two processes subequal in length) to markedly asymmetrical . (with one lateral process longer than the other), a bipennate Sc element with a shorter anterior process, a bipennate Pa element resembling the Sc but with the unit inwardly bowed with a more prominently arched basal margin, and a digyrate Pb element with two lateral processes directed in opposite directions distally.
Chirognathus cliefdenensis Zhen and Webby, 1995, from the Cliefden Caves Limestone Subgroup of central New South Wales, differs from C. duodactylus in having distinctive blade-like P elements with high processes bearing closely spaced, basally confluent denticles (Zhen and Webby 1995, pl. 2, figs 13-16).
Chirognathus tricostatus sp. nov. Figs 4-5
Synonymy
Chirognathus monodactyla Branson and Mehl; Burrett, 1979, pp. 31-32.
Tasmanognathus careyi Burrett, 1979, p. 33-35, partim, only pl. 1, fig. 12.
Derivation of name
Latin ¢ri- (three) and costatus (ribbed) referring to the distinctive character, the tricostate cusp of the Sb, Sc and Sd elements, of this Tasmanian species.
Material
71 specimens from eight samples (see Table 1). Holotype: AMF.136496, YYF5, Sd element (Fig. 5A-C); paratypes: AM F.136490, C137c, Sa element (Fig. 4A-C); AM F.136491, JRC 2, Sa element (Fig.
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AD); AM F.136492, YYF5, Sb element (Fig. 4E); AM F.136493, C137c, Sb element (Fig. 4F); AM F.136494 (=UTG96872: Burrett 1979, pl. 1, fig. 12; originally designated as one of the syntypes of T. careyi), Sb element (Fig. 4G-H); AM F.136495, YYF5, Sc element (Fig. 41-J); AM F.136497, C137c, Sd element (Fig. SD-E); AM F.136498, C137c, Sd element (Fig. 5F-G); AM F.136499, JRC 2, Pa? element (Fig. 5H); AM F.136500 (=UTG96866), JRC 2, Pa element (Fig. 51); AM F.136501, JRC 2, Pa element (Fig. 5J- K); AM F.136502, YYF1, Pb element (Fig. SL-N); AM F.136503, YYF4, Pb element (Fig. 5O).
Diagnosis
A species of Chirognathus with a seximembrate (possibly septimembrate) ramiform-ramiform apparatus including alate Sa, modified digyrate Sb and Sd, modified bipennate Sc, bipennate Pa and digyrate Pb elements; all elements with long, peg-like denticles, and a shallow, open basal cavity, typically preserved without attachment of a basal funnel.
Description
Sa element symmetrical or nearly symmetrical, with a prominent cusp and a denticulate lateral process on each side (Fig. 4A-D); cusp large, straight, antero-posteriorly compressed, with broadly convex anterior and posterior faces and sharply costate lateral margins; lateral processes extending laterally and bearing three or more denticles of variable sizes, which are also antero-posteriorly compressed; basal cavity flared anteriorly and posteriorly with basal margin nearly straight or slightly arched in posterior or anterior view (Fig. 4A, D).
Sb element (Fig. 4E-H) like Sa, but asymmetrical with outer lateral process slightly curved posteriorly and with a short, but prominent costa developed on the basal part of the anterior face (Fig. 4E, H); outer lateral process slightly curved posteriorly and also with basal margin twisted posteriorly and upper margin anteriorly (Fig. 4G); basal cavity shallow, flared anteriorly and posteriorly and extending distally as a narrow and shallow groove underneath each process (Fig. 4F).
Sc element modified bipennate, strongly asym- metrical with denticulate anterior and posterior processes and a strong costa on the outer lateral face (Fig. 4I-J); both processes extending straight or slightly curved inward; anterior process bearing three or more denticles with the distal denticle (away from the cusp) larger than the other denticles; posterior process bearing two or more denticles with the distal one (away from the cusp) larger than the other denticle; larger denticle on the posterior or anterior
Sl
LATE ORDOVICIAN CONODONTS FROM TASMANIA
Figure 4. Chirognathus tricostatus sp. nov. A-D, Sa element; A-C, AM F.136490, paratype, C137c, A, anterior view (LY 138-020), B, basal view ([Y138-021), C, posterior view (1Y142-023); D, AM F.136491, paratype, JRC 2, anterior view (IY 142-002). E-H, Sb element; E, AM F.136492, paratype, YYF5, an- terior view (IY135-039); F, AM F.136493, paratype, C137c, posterior view (LY138-022); G-H, AM F.136494=UTG96872 (Burrett 1979, pl. 1, fig. 12; originally designated as one of the syntypes of T. careyi), paratype, JRC 2, G, posterior view (IY141-018), H, anterior view (1Y141-019). I-J, Sc element, AM F.136495, paratype, YYF5, I, upper-inner lateral view (1Y 135-035), J, upper-outer lateral view (LY 135-
036). Scale bars 100 um.
process being as wide as the cusp in the lateral view, but more strongly compressed laterally than the cusp; outer lateral costa prominent, forming a ridge-like process near the base (Fig. 4J).
Sd element modified digyrate, strongly asym- metrical with a robust cusp, a denticulate lateral process on each side and a blade-like costa on the anterior face (Fig. 5A-G); cusp tricostate with a sharp costa along the lateral margins and on the broadly convex anterior face, and a less convex posterior face; anterior costa more strongly developed than that in the Sb element, and extending to near the tip of the cusp, and basally often developed into a short, blade-
a2
like process (Fig. 5C-D, G); lateral processes distally curved posteriorly bearing three or more denticles of variable sizes; basal cavity more open and strongly flared posteriorly than that of the Sb element, forming a strongly arched basal margin in posterior view (Fig. SF).
Pa element bipennate with a prominent cusp and denticulate anterior and posterior processes (Fig. 5H- K); cusp suberect, laterally compressed with sharply costate anterior and posterior margins and broadly convex lateral faces (Fig. 5H-J); both anterior and posterior processes bearing three or more denticles of variable sizes, which are also laterally compressed;
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Figure 5. Chirognathus tricostatus sp. nov. A-G, Sd element; A-C, AM F.136496, Holotype, YYF5, A, an- terior view (LY 135-034), B, posterior view (LY 142-028), C, upper view (LY135-033); D-E, AM F.136497, paratype, C137c, D, anterior view (IY 142-025), E, posterior view (LY 138-027); F-G, AM F.136498, para- type, C137c, F, posterior view (1Y 138-024), G, anterior view (LY 142-026). H, Pa? element; AM F.136499, paratype, JRC 2, outer lateral view ([Y142-018); I-K, Pa element, I, AM F.136500 =UTG96866, para- type, JRC 2, outer lateral view (TY141-026); J-K, AM F.136501, paratype, JRC 2, J, inner lateral view (1Y142-020), K, basal view, close up showing the zone of recessive basal margin (LY 142-022). L-O, Pb element; L-N, AM F.136502, paratype, YYF1, L, posterior view (LY 136-30), M, upper view (IY 136029), N, anterior view (LY 142-029); O, AM F.136503, paratype, YYF4, basal-posterior view (LY 135-026). Scale bars 100 um.
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LATE ORDOVICIAN CONODONTS FROM TASMANIA
Fig. 6. A-I, Panderodus gracilis (Branson and Mehl, 1933). A, falciform, AM F.136504, JRC 2, outer-lat- eral view (LY 139-033). B-C, truncatiform element, AM F.136505, JRC 2, B, posterior view (LY139-026); C, inner-lateral view (LY 139-024). D-G, graciliform element; D-F, AM F.136506, JRC 2, D, inner-lateral view (1Y139-017); E, outer-basal view of the basal part ([Y139-022); F, outer-lateral view (LY139-020); G, AM F.136507, JRC 2, outer-lateral view ([Y139-023). H-I, falciform element; H, AM F.136508, JRC 2, inner-lateral view ([Y139-035); I, AM F.136509, YYF2, outer-lateral view ([Y140-25). J-N, Protopan- derodus? nogamii (Lee, 1975). J, Sb element, AM F.136510, YYF4, outer-lateral view (1Y 136-027). K-N, Pa element; K-L, AM F.136511, YYF4, K, outer-lateral view (1Y 136-025), L, outer lateral view, closer up showing the furrow weaken and disappeared before researching basal margin ([Y136-026). M-N, AM F.136512, YYF3, M, outer-lateral view ([Y 140-021), N, basal view (1Y140-019). Scale bars 100 1m unless otherwise indicated.
anterior process typically slightly curved inward and extending downward forming a gently arched basal margin in lateral view (Fig. 5I-J); basal cavity shallow and open, often with zone of recessive basal margin preserved (Fig. 5K).
Pb element digyrate with a prominent cusp and denticulate lateral process on each side (Fig. 5L-O); cusp curved posteriorly with costate lateral margins; lateral processes bearing four or more denticles of variable sizes; basal cavity shallow and open with
54
gently arched basal margins in anterior or posterior view (Fig. SL, N-O).
Discussion
Chirognathus tricostatus sp. noy. was initially reported by Burrett (1979) as Chirognathus monodactyla, one of the 23 form species recognized by Branson and Mehl (1933). One of the syntypes of Tasmanognathus careyi (AM F.136494 =UTG 96872) is re-assigned herein to C. tricostatus to represent
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the Sb position (Fig. 4G; also see Burrett 1979, pl. 1, fig. 12). C. tricostatus from Tasmania differs from two currently known multi-element species of Chirognathus, C. duodactylus from the Upper Ordovician (Sandbian) of North American Mid- continent faunas and C. cliefdenensis from the Upper Ordovician (Katian) of central New South Wales, in having distinctive tricostate Sb, Sc and Sd elements.
Sweet (1982, 1988) recognized the M element for the type species, C. duodactylus. A comparable element has also recognized in the Tasmanian material of C. tricostatus, but has been assigned to the Sd position to form a symmetry transitional series with other S elements. One of the illustrated specimens of the Pa element (Fig. 5H) shows a nearly straight basal margin and posteriorly curved cusp, and may possibly represent the M element of this species. However, as only one specimen is available in the current material, it is tentatively assigned to the Pa element.
Genus PHRAGMODUS Branson and Mehl, 1933
Type species Phragmodus primus Branson and Mehl, 1933.
Phragmodus undatus Branson and Mehl, 1933 Figs 7-8
Synonymy
Phragmodus undatus Branson and Mehl, 1933, p. 115-116, pl. 8, figs 22-26; Zhen and Webby, 1995, p. 284, pl. 4, fig. 5; Leslie and Bergstrém, 1995, p. 970-973, fig. 4.1-4.14 (cum syn.); Zhen et al., 1999, p. 90, fig. 9.1- 9.5 (cum syn.); Zhen et al., 2003a, fig. 6N, O; Pyle and Barnes, 2002, figs 14.11-14.12, 15.31- 15.32; Percival et al., 2006, fig. 4A-E.
Material 116 specimens from six samples (see Table 1).
Description
M element makellate, geniculate coniform with a robust cusp and a short base triangular in outline (Fig. 7A-B); cusp strongly antero-posteriorly compressed forming sharp lateral edges and broad anterior and posterior faces; inner-lateral corner triangular in outline, and outer-lateral proto-process short with a gently arched upper margin; basal cavity shallow with weakly wavy basal margins.
S elements ramiform bearing a long multi- denticulate posterior process with one or two enlarged denticles, but none of the Tasmanian specimens
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have the posterior process completely preserved. Sa element symmetrical or nearly symmetrical with a prominent costa on each side (Fig. 7C-D); posterior process long with one denticle (typically the third or fourth from the cusp) about twice as wide as the adjacent denticles, and larger and longer than the cusp; in some specimens a costa also developed on each side of the larger denticle (Fig. 7D); basal cavity shallow with strongly arched basal margins; anterior (or antero-inner lateral) costa typically only weakly developed (Fig. 7D). Sb element modified quadriramate, like Sa but asymmetrical with the sharp costate anterior margin curved inward (Fig. 7F-G). Sc element modified bipennate, like Sb but strongly asymmetrical with a sharply costate anterior margin curved inward and with smooth inner and outer lateral faces (Fig. 7H-L). Sd element tertiopedate, like Sb, but with a broad anterior face and with one of the larger denticles on the posterior process curved inward and the other outward (Fig. 8A-C).
Pa element pastinate with long denticulate posterior and inner lateral processes, and a suberect cusp (Fig. 8D-G); cusp laterally compressed with sharply costate anterior and posterior margins, outer lateral face more convex; posterior process long, bearing six or more denticles; inner lateral process shorter, bearing five or more denticles and strongly bending anteriorly forming an angle of nearly 180 degree with the posterior process (Fig. 8E, G); costate anterior margin extending downward and not forming a prominent anterior process (Fig. 8D); basal cavity shallow, forming a wide and open groove along the posterior and inner lateral processes, and flared anteriorly and inner laterally (Fig. 8G). Pb element pastinate, like Pa but with a more robust cusp and less anteriorly curved inner lateral process (Fig. 8H-I).
Discussion
Leslie and Bergstr6m (1995) suggested a seximembrate apparatus for P undatus, including adenticulate makellate M, alate Sa, tertiopedate Sb, bipennate Sc, pastinate Pa and Pb elements. All six elements have been recovered from the Tasmanian samples (Figs 7-8); they are identical with those described and illustrated by Leslie and Bergstrém (1995, fig. 4) from the Joachim Dolomite and Kings Lake Limestone of Missouri, except that an additional tertiopedate element was recognized in the Tasmanian material (Fig. 8A-C). This latter element is similar to the Sb element, but has the cusp and the larger denticles on the posterior process strongly twisted towards different sides in respect to the antero- posterior axis. It is assigned herein to represent the Sd position.
5
LATE ORDOVICIAN CONODONTS FROM TASMANIA
Fig. 7. Phragmodus undatus Branson and Mehl, 1933. A-B, M element; A, AM F.136513, YYF4, posterior view (1Y136-005); B, AM F.136514, YYF4, anterior view (1Y136-006). C-D, Sa element, AM F.136515, C137c, C, basal view (LY 138-014); D, lateral view (LY138-015). E-G, Sb element; E, AM F.136516, YYF4, outer-lateral view (LY 136-015); F, AM F.136517, YYF4, inner-lateral view (LY 136-014), G, AM F.136518, YYF4, inner-lateral view (1Y136-016). H-L, Sc element; H, AM F.136519, YYF4, inner-lateral view (LY 136-013); I-J, AM F.136520, YYF4, I, outer-lateral view (LY136-009), J, inner-lateral view (1Y136- 010); K-L, AM F.136521, C137c, K, basal view (LY 138-016), L, outer-lateral view (1Y 138-017). Scale bars 100 um.
56 Proc. Linn. Soc. N.S.W., 131, 2010
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Fig. 8. Phragmodus undatus Branson and Mehl, 1933. A-C, Sd element; AM F.136522, JRC 2, A, upper view (IY 138-028), B, outer-lateral view (IY 138-029), C, posterior view (ITY 138-030). D-G, Pa element; D- E, AM F.136523, YYF4, D, outer-lateral view ([Y136-001), E, basal view ([Y136-011); F-G, AM F.136524, YYF4, F, inner-lateral view (LY136-003), G, basal view ([Y136-012). H-I, Pb element; H, AM F.136525, YYF4, outer-lateral view (1Y¥136-004); I, AM F.136526, YYF4, antero-outer lateral view (1Y136-017).
Scale bars 100 um.
Genus PROTOPANDERODUS Lindstrém, 1971 Synonymy Scolopodus nogamii Lee 1975, p. 179, pl. 2, fig. 13.
?Panderodus nogamii (Lee); Cantrill and Burrett 2004, p. 410, pl. 1, figs 1-16.
Panderodus nogamii (Lee); Zhang et al. 2004, p. 16, pl. 5, figs 1-5.
Protopanderodus nogamii (Lee); Watson 1988: p. 124, pl. 3, figs 1, 6; Zhen et al. 2003b, p. 207-
Type species Acontiodus rectus Lindstrém, 1955.
Protopanderodus? nogamii (Lee, 1975) Fig. 6J-N
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LATE ORDOVICIAN CONODONTS FROM TASMANIA
209, fig. 23A-P, ?Q (cum syn.); Zhen and Percival 2004a, p. 104-105, fig. I8A-K (cum syn.).
Protopanderodus? nogamii (Lee); Zhen and Percival 2004b, p. 170-172, fig. 11P, Q (cum syn.).
Material 69 specimens from four samples (see Table 1).
Discussion
Recent review of this species by Cantrill and Burrett (2004) suggested a geographical distribution restricted to Gondwana and_peri-Gondwanan terranes. Morphologically P nogamii is_ rather conservative over its long stratigraphic range from the upper Floian (evae Zone, Zhen et al. 2003b) to upper Sandbian (uwndatus Zone, this study). Generic assignment of this species has been debated in the literature (see synonymy list). Most elements of this species bear a non-panderodontid furrow on each side, suggesting that it might be more closely related to Protopanderodus rather than to typical Panderodus.
Genus TASMANOGNATHUS Burrett, 1979
Type species Tasmanognathus careyi Burrett, 1979.
Diagnosis
Septimembrate apparatus with a ramiform- pectiniform apparatus structure including makellate M, ramiform S (including alate Sa with a denticulate lateral process on each side, digyrate Sb, bipennate or modified bipennate Sc, and tertiopedate Sd), carminate Pa, and angulate Pb (some species with an additional modified angulate or pastinate Pb2) elements.
Discussion
Following Burrett’s (1979, p. 32) original view that Zasmanognathus might be closely related to Rhipidognathus, Aldridge and Smith (1993) doubtfully included it in the Rhipidognathidae. Affinities with other genera remain conjectural, although greatest similarities appear to be with Yaoxianognathus (see discussion below).
Tasmanognathus was established on a single species, 7. careyi Burrett, 1979 from the Lower Member of the Benjamin Limestone in the Florentine Valley and Everlasting Hills of central Tasmania. Subsequently, Zasmanognathus has been reported from the mid Darriwilian to upper Katian of eastern Australia, North China (An et al. 1985, An and Zheng 1990, Pei and Cai 1987), Qinling Mountains in the Kunlun-Qinling Region (Pei and Cai 1987), Tarim
58
Basin (Zhao et al. 2000; Jing et al. 2007), South Korea (Lee 1982; An and Zheng 1990), ?Siberia and northeastern Russia (Domoulin et al. 2002), and possibly North America (where it was referred to as Yaoxianognathus abruptus). It is represented by nine named species and several additional unnamed forms, the latter included herein in 7Zasmanognathus although some are poorly known or inadequately documented. Following is a brief review of the known species (with our interpretation of element notations in parentheses):
Tasmanognathus careyi Burrett, 1979 from the Lower Limestone Member of the Benjamin Limestone in the Florentine Valley and Everlasting Hills of central Tasmania; a seximembrate apparatus was originally recognized, but based on re-examination of original topotypes and additional new material, it has been revised herein as having an septimembrate apparatus (including M, Sa, Sb, Sc, Sd, Pa, and Pb elements).
Badoudus badouensis Zhang in An et al., 1983 from the Fengfeng Formation (Sandbian) of Handan, Hebei Province in North China (considered by An et al. 1985, p. 102, to represent a species of Tasmanognathus); this is a poorly defined form species with only two specimens illustrated (An et al. 1983, pl. 25, figs 5, 6, text-fig. 12.17), both of which are carminate, bearing an indistinctive cusp and a long denticulate anterior process and a short denticulate posterior process. This element is comparable with the Pa element of Zasmanognathus defined herein.
Tasmanognathus borealis An in An et al., 1985 from the upper part of the Yaoxian Formation (late Sandbian) of Yaozhou District (formerly Yaoxian) of Tongchuan City, Shaanxi Province in North China; originally defined as having a quinquimembrate apparatus, including trichonodelliform (= Sa element; see An et al. 1985, pl. 1, fig. 20), zygognathiform (© Sbyelement: see An”etialsalOSse pkely fis. 13) cordylodiform (= Sc element; see An et al. 1985, pl. 1, fig. 15), ozarkodiniform (= Pa element; see An et al. 1985, pl. 1, fig. 14), and prioniodiniform (= Pb element; see An et al. 1985, pl. 1, fig. 16).
Tasmanognathus gracilis An in An et al., 1985 from the upper part of the Yaoxian Formation (late Sandbian) of Yaozhou District (formerly Yaoxian) of Tongchuan City, Shaanxi Province in North China; originally defined as having a seximembrate apparatus, including cyrtoniodiform (= M element; see An et al. 1985, pl. 1, fig. 8), trichonodelliform (= Sa element; see An et al. 1985, pl. 1, fig. 12), ligonodiniform (= Sb element; see An et al. 1985, pl. 1, fig. 11), cordylodiform (= Sc element; see An et al. 1985, pl. 1, fig. 10), ozarkodiniform (= Pa element:
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see An et al. 1985, pl. 1, fig. 7), and prioniodiniform (= Pb element; see An et al. 1985, pl. 1, fig. 9).
Tasmanognathus multidentatus An in An and Zheng, 1990 (p. 20, 95, text-fig. 9, pl. 11, fig. 4); the only figured specimen (pl. 11, fig. 4) is a Pa element from the Yaoxian Formation of Yaozhou District (formerly Yaoxian) of Tongchuan City, Shaanxi Province in North China, which is identical with the Pa element of 7. borealis An in An et al., 1985. In fact, the figured Pa element (pl. 11, fig. 4) of 7 multidentatus and the holotype and a figured paratype of T: borealis (An et al. 1985, pl. 1, figs 13, 16) were recovered from the same sample (Tpl3y2). It is unclear why An and Zheng (1990) tried to replace 7. borealis with T: multidentatus. However, as the latter is anomem nudum, T. borealis remains the valid name for this Yaoxian species.
Tasmanognathus planatus Pei in Pei and Cai, 1987 from the Sigang Formation of Xichuan and Neixiang Counties, Henan Province in the Qinling Mountains . (Pei and Cai 1987; Chen et al. 1995; Wang et al. 1996); the type material was represented by Pa (Pei and Cai 1987, pl. 13, fig. 12), Pb (Pei and Cai 1987, pl. 13, figs 8, 713), and Sb (Pei and Cai 1987, pl. 13, fig. 9) elements.
Tasmanognathus shichuanheensis An in An et al., 1985 from the lower part of the Yaoxian Formation (mid Sandbian) of Yaozhou District (formerly Yaoxian) of Tongchuan City, Shaanxi Province in North China; originally defined as having a seximembrate apparatus, including cyrtoniodiform GuvMmclementsisceAnwetale 1985s) pla tie. 3): trichonodelliform (= Sa element; see An et al. 1985, pl. 1, fig. 4), ligonodiniform (= Sb element; see An et al. 1985, pl. 1, fig. 1), cordylodiform (= Sc element; see An et al. 1985, pl. 1, fig. 5), ozarkodiniform (= Pa element; see An et al. 1985, pl. 1, fig. 2), and prioniodiniform (= Pb element; see An et al. 1985, pl. 1, fig. 6).
Tasmanognathus sigangensis Pei in Pei and Cai, 1987 from the Shiyanhe Formation (late Sandbian- early Katian) of Neixiang County, Henan Province in the Qinling Mountains; a quinquimembrate species apparatus was recognized including trichnodelliform (= Sa element; Pei and Cai 1987, pl. 13, fig. 4), zygognathiform (= Sb element, Pei and Cai 1987, pl. 13, fig. 11), cordylodontiform (= Sc element; Pei and Cai 1987, pl. 13, fig. 7), prioniodiniform (= Pa element; Pei and Cai 1987, pl. 13, figs 1-2), and ozarkodontiform (= ?Pb element; Pei and Cai 1987, joll, 13}, 10g, 3),
Tasmanognathus sishuiensis Zhang in An et al. 1983 reported from the upper Fengfeng Formation (early Sandbian) of Shandong and Hebei
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provinces in North China; defined as consisting of a quinquimembrate apparatus including trichonodelliform (= Sa element, see An et al. 1983, pl. 29, figs 7, 9, 10), zygognathiform (= Sb element, see An et al. 1983, pl. 29, figs 4-6, 8, ?11), cordylodontiform (= Sc element, see An et al. 1983, pl. 29, figs 1-3), ozarkodiniform (= Pa element, see Anetal. 1983, pl. 29, figs 14-15), and prioniodiniform (= Pb element, see An et al. 1983, pl. 29, figs 12-13) elements. This species is characterized by its widely spaced peg-like denticles on the S elements.
Tasmanognathus sp. described by Pei and Cai (1987) from the Sigang and Shiyanhe formations of Neixiang County, Henan Province in the Qinling Mountains; represented by cordylodontiform (= Sc element; Pei and Cai 1987, pl. 13, figs 5-6) and prioniodiniform (= ?Pb element; Pei and Cai 1987, pl. 13, fig. 10) elements.
Tasmanognathus sp. from the Fossil Hill Limestone (early Katian) of the Cliefden Caves Limestone Subgroup, central New South Wales was only represented by the Pa element (Zhen and Webby 1995, p. 289, pl. 5, fig. 23), which showed close resemblance to the Pa element of 7: borealis from the Yaoxian Formation.
Tasmanognathus sp. cf. T. borealis An in An et al., 1985; only the Pa element known from unnamed limestone of Late Ordovician (late Sandbian) age intersected in drillcore in the Marsden district of south-central New South Wales (Percival et al. 2006).
The three species of Zasmanognathus (T. borealis, T: gracilis and T. shichuanheensis) erected by Anin An etal. (1985) from the Yaoxian Formation (Darriwilian- Sandbian) of Yaozhou District (formerly Yaoxian) of Tongchuan City, Shaanxi Province in North China exhibit similar species apparatus and closely related morphological variations of constituent elements. An et al. (1985) established two conodont zones in the Yaoxian Formation, namely the T. shichuanheensis Zone in the lower part of the formation (Bed 1 to Bed 3, see An et al. 1985, fig. 2), and the Zasmanognathus borealis-T. gracilis Zone spanning the upper part of the Yaoxian Formation (Bed 4 to Bed 8) into the basal part of the overlying Taoqupo Formation (Bed 9). An and Zheng (1990, p. 95, text-fig. 9) suggested that 7: sishuiensis from the Fengfeng Formation might be the direct ancestor of the species from the Yaoxian Formation, and indicated an inferred lineage from 7. sishuiensis to T: shichuanheensis and then to T. multidentatus (= T. borealis). They showed the morphological changes of the three species, mainly from widely spaced denticles on the processes of the S and Pb elements and a prominent cusp on the Pa
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element of 7: sishuiensis, to closely spaced denticles in the S and Pb elements and an indistinctive cusp inthe Pa element of 7. mu/tidentatus (= T. borealis). However, these species from the Yaoxian Formation and Fengfeng Formation show some detailed differences in composition of the apparatus in comparison with T. careyi from Tasmania. In particular, they seem to lack makellate M and tertiopedate Sd elements, and have a “dolabrate” Sc element with a long denticulate posterior process. Morphologically, such features support a closer relationship with Yaoxianognathus yaoxianensis An in An et al., 1985. However, these species lack hindeodellid denticles on the processes of the S elements, which was the major character that An (in An et al. 1985) employed to distinguish Yaoxianognathus from Tasmanognathus. As revision of An’s species of Zasmanognathus from the Yaoxian Formation and the Fengfeng Formation of North China is beyond the scope of the current study, they are retained in Zasmanognathus for the time being, although they show some significant differences in morphology and apparatus composition in comparison with the type species of Zasmanognathus as revised here.
Based on the concept of Yaoxianognathus employed by An (in An et al. 1985) and others (e.g. Savage 1990; Zhen et al. 1999), generic assignment of species previously included in Yaoxianognathus but which apparently lack hindeodellid denticles on the processes of the S elements, should be reconsidered. For example, Yaoxianognathus abruptus (Branson and Mehl, 1933), a North American Midcontinent species ranging across the wndatus to tenuis zones of the Mohawkian, was initially proposed as a form species based only ona carminate Pa element (Branson and Mehl, 1933, pl. 6, fig. 11) and revised by Leslie (2000, p. 1143) as having a seximembrate apparatus. It closely resembles An’s species of Tasmanognathus from North China; most importantly, none of Leslie’s illustrated S elements of ¥. abruptus (fig. 4.15-4.18) bears hindeodellid denticles that are characteristic of Yaoxianognathus, and hence we suggest this species more likely belongs to Tasmanognathus.
Similarly, S elements of Yaoxianognathus? neonychodonta Zhang, Barnes and Cooper, 2004, from the Stokes Siltstone of the Amadeus Basin in central Australia, lack hindeodellid denticles and therefore should be excluded from Yaoxianognathus. As Zhang et al. (2004) implied, this species may be more closely related to Plectodina, judging from the morphological characters of its ramiform S and pastinate Pb elements.
In comparison, the two multielement species of Yaoxianognathus from the Upper Ordovician of
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central New South Wales (¥. wrighti Savage, 1990 and Y. ani Zhen, Webby and Barnes, 1999) do exhibit well developed hindeodellid denticles on the processes of the S elements, particularly on the long posterior process of the Sc element (Savage 1990, fig. 6.7-6.12; Zhen et al. 1999, fig. 15.3-15.6, 15.9-15.12, 15.16). The apparatuses of both species include a makellate M and a modified bipennate Sc elements, which differ morphologically from corresponding elements in the T. careyi apparatus as defined herein.
Tasmanognathus careyi Burrett, 1979 Figures 9-15
Synonymy
Tasmanognathus careyi Burrett, 1979, p. 33-35, partim only text-figs 2-4, pl. 1, figs 1-7, 11, 13-19 (text-fig. 2 = Pb2 , text-fig. 3 = Pa, text-fig.4A = Sb, text-fig. 4B = Sc, text-fig. 4C, D = Sd; pl. 1, figs 1-3 = Pb2, 4-5 = Pbl, 6-7 = Pa, fig. 11 = Sc, figs 13-14= Sb, figs 15-18 = Sd, fig. 19 = Sa); non fig. 12 = C. tricostatus sp. nov., non figs 8-10, 20 =, sp. chcareyi.
? Tasmanognathus careyi Burrett; An and Zheng, LO9Os pl alalchiesy 2)
Material
297 specimens from nine samples (see Table 1).
Burrett (1979, p. 33, pl. 1, figs 1-7, 11-12, 17-18, 20) designated 11 figured specimens from sample JRC 2 as syntypes, ten of which (excluding UTG 96863 which was not able to be located for this study; figured by Burrett 1979, pl. 1, figs 17-18), and 225 additional specimens (including originally undesignated topotypes) from five samples (LLMB, C137, C98, JRC 2 and JRB, see Table 1) are available for the current study. AM F.136547 (~UTG 96851; Burrett 1979, pl. 1, fig. 6) representing a Pa element is selected herein as lectotype (Fig. 14A-B); and seven out of ten originally designated and illustrated syntypes were examined and illustrated herein as paralectotypes, including AM F.136557 (=UTG 96857, Fig. 15H; Burrett 1979, pl. 1, fig. 1), AM F.136559 (-UTG 96860; Fig. 15K; Burrett 1979, pl. 1, fig. 2), AM F.136560 (=UTG 96853, Fig. 15L; Burrett 1979, pl. 1, fig. 3), AM F.136553 (~UTG 96850, Fig. 15A-B; Burrett 1979, pl. 1, fig. 4), AM F.136554 (=UTG 96882, Fig. 15C; Burrett 1979, pl. 1, fig. 5), AM F.136548 (~UTG 96856, Fig. 14C; Burrett 1979, pl. 1, fig. 7), and AM F.136539 (~UTG 96876, Fig. 12A-C; Burrett 1979, pl. 1, fig. 11).
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Figure 9. Tasmanognathus careyi Burrett, 1979. M element; A, AM F.136527 =UTG96875, JRC 2, ante- rior view (1Y141-025). B, AM F.136528, C137c, anterior view ([Y138-011). C-D, AM F.136529, C137c, C, posterior view (1Y 138-008); D, basal view (TY138-009). E-F, AM F.136530, JRC 2, E, posterior view (LY 138-035); F, basal view (1Y138-034). G-J, AM F.136531, JRC 2, G, upper view (1Y139-007); H, pos- terior view (LY139-008); I, inner-lateral view (LY139-009); J, anterior view (1Y139-010). Scale bars 100
um.
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Figure 10. Tasmanognathus careyi Burrett, 1979. Sa element; A-B, AM F.136532 =UTG96874 (Burrett 1979, pl. 1, fig. 19), JRC 2, A, posterior view (TY 141-022), B, lateral view (LY141-021); C-E, AM F.136533, YYF5, C, anterior view (1Y140-004), D, posterior view ([Y140-003), E, upper-posterior view (1Y140- 001); F-I, AM F.136534, YYF4, F, lateral view (TY135-019), G, posterior view (1Y135-020), H, anterior view (LY135-018), I, upper view, close up showing the cross section of the cusp ([Y 135-022). Scale bars 100 um.
UTG 96877, previously designated as a syntype (Burrett 1979, pl. 1, fig. 20) is excluded from this species and re-assigned to T. sp. cf. careyi representing the Sb position (AM F.136567, Fig. 16G herein). Another previously designated syntype UTG 96872 (Burrett 1979, pl. 1, fig. 12) is also excluded from this species and re-assigned to Chirognathus tricostatus sp. nov. where it represents the Sb position (AM F.136494, Fig. 4G-H herein).
Diagnosis
Septimembrate apparatus with a ramiform- pectiniform structure including makellate M, alate Sa, digyrate Sb, bipennate Sc, tertiopedate Sd, carminate Pa, angulate Pbl, and pastinate Pb2 elements. S elements with a robust cusp, an open and shallow basal cavity, and long closely-spaced denticles on the processes; Pa element with a longer anterior process,
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a nearly straight basal margin and a cusp varying from prominently larger (juvenile) than adjacent denticles to rather indistinctive in size (when mature). Pb1 element with a robust cusp, and a strongly curved basal margin. Pb2 element with a short adenticulate outer lateral process, long denticulate anterior and posterior processes, and a strongly laterally flared base.
Description
M element makellate with a denticulate inner- lateral process bearing three to five pointed denticles (Fig. 9), and a shorter, typically adenticulate outer lateral process (Fig. 9A-C, H); cusp robust, antero- posteriorly compressed (Fig. 9G), with a sharp costa along the inner-lateral and outer lateral margins (Fig. 9G-I), and distally curved posteriorly (Fig. 9C-D, G); denticles on the inner lateral process also antero-
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Figure 11. Jasmanognathus careyi Burrett, 1979. Sb element; A, AM F.136535 =UTG96873, posterior view (1Y141-027); B-D, AM F.136536, C98, B, basal view ([Y137-039), C, anterior view (1Y137-037), D, basal-posterior view (LY137-038); E, AM F.136537, C98, outer-anterior view (LY 137-034); F-H, AM F.136538 =UTG96898 (Burrett, 1979, fig. 4A), C98, F, anterior view (TY137-031), G, upper view (IY 137- 032), H, upper-posterior view (LY 137-030). Scale bars 100 um.
posteriorly compressed, with a sharp costa along the inner-lateral and outer-lateral margins (Fig. 9C, G- H); basal cavity shallow and open, tapering towards distal ends of the processes and flaring posteriorly (Fig. 9D, F), and often with weakly developed zone of recessive basal margins (Fig. 9F); anterior portion of basal margin nearly straight (Fig. 9B, J), but posterior portion weakly curved (Fig. 9C, E, H).
Sa element alate (Fig. 10), symmetrical with a robust cusp, a prominent tongue-like anticusp, and a long denticulate lateral process on each side; cusp proclined, subquadrate in cross section (Fig. 10E, I), with a sharp costa on each side (Fig. 10A-B) and often a weak costa along the posterior margin (Fig. 10D-E), but some specimens with a broad posterior face (Fig. 10G) or with a broad carina developed (Fig. 10A); broad anterior face bearing a shallow but prominent mid groove and a broad carina on each side (Fig. 10C, H); cusp extended downward to form a downward extending tongue-like anticusp (Fig. 10A-D, H); lateral process long, bearing up to ten or more closely spaced denticles (Fig. 10C-D, H), which
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are compressed antero-posteriorly; basal cavity open and shallow, flared posteriorly; basal margin arched in posterior view (Fig. 10D).
Sb element digyrate, asymmetrical, with a robust cusp, long denticulate process on each side, and a prominent downwardly extending tongue-like anticusp (Fig. 11); cusp suberect, slightly curved inward (Fig. 10A), with a more strongly convex anterior face, and a sharp costa on each side (Fig. 11G-H); outer lateral process shorter, bearing three or more denticles (Fig. 11A, D, G); inner lateral process longer, bearing five or more peg-like denticles (Fig. 11C, F), and more strongly curved posteriorly (Fig. 11B, G), forming an angle of about 100-110 degrees between the two processes in the upper or basal view (Fig. 11B, G).
Sc element bipennate, asymmetrical with a robust cusp, a long denticulate posterior process, and a short denticulate anterior process (Fig. 12); cusp suberect basally and reclined distally (Fig. 12A, F, H) with a more convex outer lateral face, and laterally compressed with a sharp costa forming anterior
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Figure 12. Tasmanognathus careyi Burrett, 1979. Sc element; A-C, AM F.136539 =UTG96876 (Burrett 1979, pl. 1, fig. 11), JRC 2, paralectotype, A, inner lateral view (1Y141-020), B, basal view (LY 141-016), C, outer lateral view (LY141-015); D-E, AM F.136540, YYF5, D, inner-basal view (LY 135-041); E, inner-lat- eral view (LY 135-040); F-G, AM F.136541, JRC 2, F, inner lateral view (LY 139-028); G, inner-basal view (TY 139-027); H, AM F.136542, YYFS5, inner-lateral view (1Y140-009); I-J, AM F.136543 =UTG 96899 (Burrett, 1979, fig. 4B), C98, I, inner-lateral view ([Y137-029), J, basal view (LY137-027). Scale bars 100 pm.
and posterior margins (Fig. 12F-I); anterior margin curved inward (Fig. 12D-I); posterior process bearing three or more (up to seven) denticles, which are laterally compressed and posteriorly reclined (Fig. 11A, C, I); anterior process with upper margin curved inwards, and extending downwards bearing two to four small denticles (Fig. 12D-H); basal cavity open and shallow, slightly flared inwards (Fig. 12B, D, G), some specimens with basal funnel attached (Fig. 12I- I:
Sd element tertiopedate, weakly asymmetrical to nearly symmetrical with a robust cusp, a prominent anticusp, a denticulate posterior process and a denticulate lateral process on each side (Fig. 13); cusp with a broad anterior face (Fig. 13B-C), and with a prominent costa along the posterior margin and on each lateral side (Fig. 13D, G); anticup short and downward extending (Fig. 13C-D); posterior process
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long and straight, broken in most specimens, in one of the examined specimens bearing ten denticles (Fig. 13A); lateral process bearing four or more denticles (Fig. 13B-D); basal cavity open, T-shaped in basal view (Fig. 13B).
Pa element carminate (Fig. 14), laterally compressed and blade-like, with a small cusp, and with the anterior and posterior processes bearing basally confluent denticles; cusp erect (smaller specimens, Fig. 14C, E) to slightly inclined (larger specimens, Fig. 14A, D), typically larger and higher than adjacent denticles (Fig. 14C, E, F), but less distinctive in the larger specimens (Fig. 14 A, H); two processes of unequal length, anterior process longer and higher, bearing five to eight closely- spaced denticles; posterior process lower and shorter, bearing two to six denticles, with distal end slightly . bent downward (Fig. 14A, D); juvenile specimens
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Figure 13. Tasmanognathus careyi Burrett, 1979. Sd element; A, AM F.136544 =UTG96902 (Burrett, 1979, pl. 1, figs 15-16), LLMB, upper view (I1Y137-033); B-E, AM F.136545 =UTG96900 (Burrett, 1979, fig. 4C-D), B, basal view (1Y137-023), C, anterior view (LY 137-022), D, lateral view ([Y127-024), E, close up showing fine striae on the surface of the cusp (1Y137-026); F-G, AM F.136546, YYF1, G, lateral view (LY140-015), F, basal-posterior view (LY 140-014). Scale bars 100 um.
Figure 14. Tasmanognathus careyi Burrett, 1979. Pa element; A-B, AM F.136547 -UTG96851 (Burrett 1979, pl. 1, fig. 6), lectotype, JRC 2, A, outer lateral view ([Y141-002), B, basal-inner lateral view (1Y141- 003); C, AM F.136548 =UTG96856 (Burrett 1979, pl. 1, fig. 7), paralectotype, JRC 2, outer lateral view (1Y 141-004); D, AM F.136549 =UTG96893a (Burrett, 1979, fig. 3), LLM (B), inner-lateral view (LY 137- 001); E, AM F.136550 =UTG96893b (Burrett, 1979, fig. 3), LLM (B), outer-lateral view (LY 137-003); F- G, AM F.136551, C98, F, outer-lateral view (1Y137-005), G, upper view (LY 137-006); H-I, AM F.136552, JRC 2, H, inner-lateral view (LY 137-010), I, basal view (LY137-009). Scale bars 100 pm.
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Figure 15. Tasmanognathus careyi Burrett, 1979. A-G, Pb1 element; A-B, AM F.136553 =UTG96850 (Burrett 1979, pl. 1, fig. 4), paralectytype, JRC 2, A, basal-outer lateral view (1Y141-007), B, outer lat- eral view (1Y141-006); C, AM F.136554 =UTG96882 (Burrett 1979, pl. 1, fig. 5), paralectotype, JRC 2, inner lateral view (1Y 141-008); D-E, AM F.136555, C98, D, basal view (LY137-018), E, inner lateral view (TY 137-019); F-G, AM F.136556, C98, F, outer-lateral view (LY 137-021), G, basal view (1Y 137-020). H-M, Pb2 element; H, AM F.136557 =UTG96857 (Burrett 1979, pl. 1, fig. 1), paralectotype, JRC 2, outer lat- eral view (TY 141-009); I-J, AM F.136558, JRC 2, I, inner-lateral view (1Y137-040), J, basal view (TY 137- 041); K, AM F.136559 =UTG96860 (Burrett 1979, pl. 1, fig. 2), paralectotype, JRC 2, outer lateral view (TY 141-011); L, AM F.136560 =UTG96853 (Burrett 1979, pl. 1, fig. 3), paralectotype, JRC 2, outer lateral view (1Y141-012); M, AM F.136561, JRC 2, outer-lateral view (1Y137-014). Scale bars 100 um.
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exhibiting a prominently lower and shorter posterior process with two to four less closely-spaced denticles (Fig. 14C, E); basal cavity shallow and open, flared laterally and extended toward distal end of the processes as a tapering shallow groove (Fig. 141); basal margin nearly straight to slightly arched in lateral view (Fig. 14A, C-E, F).
Pbl1 element angulate (Fig. 15A-G), laterally compressed and blade-like, with a robust cusp, and denticulate anterior and posterior processes; cusp strongly compressed laterally, more convex outer laterally, suberect and slightly curved inwards with sharp anterior and posterior margins; two processes typically sub-equal in length (Fig. 15E) or with slightly longer posterior process (Fig. 15B, C), bearing three to six short, laterally compressed and basally confluent denticles; anterior process extending downward forming an angle of about 100- 120 degrees between the two processes in lateral view (Fig. 15B, F); basal cavity shallow and open, laterally
. flared and extended as a shallow groove underneath each process (Fig. 15D, G).
Pb2 element pastinate (likely a variant of the Pb1 element), with a robust cusp, long denticulate anterior and posterior processes, and a short adenticulate outer lateral process (Fig. 15H-M); cusp suberect (Fig. 15H), laterally compressed, with a broad smooth inner lateral face, and sharp anterior and posterior margins, outer lateral face smooth (Fig. 15L-M) or with a mid costa (Fig. 15H, K); anterior process typically longer, bearing up to seven or more denticles, which are typically closely spaced with confluent bases (Fig. 151, M); most specimens with posterior process broken, bearing up to five denticles (Fig. 15L); outer lateral process typically represented by a prominent tongue-like basal extension (Fig. 15J-M), or as a short adenticulate process (Fig. 15H); basal cavity shallow, outer laterally flared more strongly, and tapering as a shallow groove to the distal end of anterior and posterior processes (Fig. 15I-J).
Discussion
One originally designated syntype (AM F.136567 =UTG96877 Fig. 16G; also see Burrett 1979, pl. 1, fig. 20) and an additional figured specimen (AM F.136562 =UTG96904, Fig. 16A; also see Burrett 1979, pl. 1, figs 8-10) of T. careyi are excluded from this species and re-assigned to represent the Sb and Pb2 elements of T. sp. cf. careyi, as they exhibit more widely spaced denticles on the processes.
The original definition of the S element given by Burrett (1979) is more or less followed herein, except that his Sal element (Burrett 1979, fig. 4C-D) is now assigned to the Sd position (Fig. 13), the digyrate
Proc. Linn. Soc. N.S:W., 131, 2010
element with a longer inner lateral process (Burrett 1979, fig. 4A, referred to as Sc) to the Sb position (Fig. 11), and the bipennate element with a shorter downwardly extended and inner laterally curved anterior process (Burrett 1979, fig. 4B, referred to as Sb) to the Sc position (Fig. 12). Burrett (1978, p. 34) further recognized an Sa2 element with the cusp exhibiting a subquadrate cross section, but illustrated it as Sa (Burrett 1979, pl. 1, fig. 19; also Fig. 10A- B herein). This symmetrical or nearly symmetrical element (Fig. 10) is confirmed as occupying the Sa position. The makellate M element (Fig. 9) described herein was not recognized in Burrett’s original description of TZ careyi. Specimens originally included in the Pa element by Burrett (1979) show two morphotypes, which are defined herein to represent the Pb! (Burrett 1979, pl. 1, figs 4-5; Fig. 1SA-G) and Pb2 (Burrett 1979, pl. 1, figs 1-3; Fig. 15H-M) elements. They can be easily differentiated from each other by having a short tongue-like outer lateral process, a costa on the outer lateral face or a short adenticulate outer lateral process in the Pb2 element (Fig. 15H-M).
Burrett (1979, pp. 33-34) discussed the considerable ontogenetic variations among the P elements, in particular the posterior process of the Pa element (referred to as the Pb element by Burrett, 1979, see p. 33, fig. 3) and the anterior process of the Pb2 element (assigned to part of the Pa element by Burrett, 1979, see p. 33, fig. 2). Juveniles of the Pa element have a larger cusp and a lower posterior process with less closely spaced denticles (Fig. 14C, E; Burrett 1979, fig. 3). It cannot presently be established whether the distinctions between the Pb1 and Pb2 elements, and within the Pa elements, represent ecophenotypic variations, or whether they reflect a high degree of morphological plasticity.
T. careyi has been widely reported from North China (Zhao et al. 1984; Wang and Luo 1984; Pei and Cai 1987; An and Zheng 1990). However, judging from the illustrations of these specimens, none can be confidently assigned to the Tasmanian species, except for one specimen figured by An and Zheng (pl. 11, fig. 2) from the lower part of the Yaoxian Formation in the Ordos Basin of Shaanxi Province that is comparable with the Pa element of 7: careyi. Pa elements of 7: borealis (An in An et al. 1985, pl. 1, fig. 14) and 7. multidentatus (An and Zheng 1990, pl. 11, fig. 4; = T. borealis), also from the Yaoxian Formation of the Ordos Basin, similarly have an indistinct cusp that is nearly the same size as adjacent denticles, but the outline of these two illustrated specimens is shorter and higher in comparison with the Pa element of 7. careyi (Fig. 14).
67
LATE ORDOVICIAN CONODONTS FROM TASMANIA
Figure 16. Tasmanognathus sp. cf. careyi Burrett, 1979. A, Pb2 element, AM F.136562 =UTG96904 (Bur- rett, 1979, pl. 1, figs 8-10), LLM (B), outer-lateral view (1Y 137-007). B, M element, AM F.136563, YYF1, posterior view ([Y136-032). C-F, Sa element; C-D, AM F.136564, JRC 2, C, Posterior view (LY138-006); D, postero-basal view (LY 138-005); E, AM F.136565, JRC 2, posterior view (LY 139-029); F, AM F.136566, YYF1, anterior view (1Y140-010). G, Sb element, AM F.136567 =UTG96877 (syntype of T. careyi; Bur- rett 1979, pl. 1, fig. 20), JRC 2, posterior view (1Y141-024). H-I, Sd element, AM F.136568, YYF4, H, posterior view (IY 135-023), I, upper view (LY135-024); J-K, Sc element, AM F.136569, YYF1, J, inner lateral view (1Y140-013), K, outer lateral view (1Y140-012). Scale bars 100 um.
Tasmanognathus sp. cf. T. careyi Burrett, 1979 Figures 16-17
Synonymy
Tasmanognathus careyi Burrett, 1979, p. 33-35, partim only pl. 1, figs 8-10 (= Pb2 element), fig. 20 (= Sb element).
Material
34 specimens from four samples in the Settlement Road section of Florentine Valley area (see Table 1).
68
Diagnosis
A species of Zasmanognathus septimembrate apparatus, including makellate M, alate Sa, digyrate Sb, bipennate Sc, digyrate? (modified tertiopedate) Sd, carminate Pa, angulate? (bipennate) Pb1, and pastinate Pb2 elements; elements robust and large in size bearing a prominent cusp ornamented with fine striae, and small widely spaced denticles on the processes of M, S, Pb1 and Pb2 elements; most elements with basal funnel attached.
having an
Proc. Linn. Soe. N.SOW.a3 1, 2010
Y.Y. ZHEN, C.F. BURRETT, I.G. PERCIVAL AND B.Y. LIN
Fig. 17. Tasmanognathus sp. cf. T. careyi Burrett, 1979. A-H, Pa element; A-C, AM F.136570, YYF4, A, inner-lateral view (LY135-005), B, upper view (LY135-003), C, outer-lateral view, close up showing fine surface striae (1Y135-007); D-F, AM F.136571, YYF4, D, outer-lateral view (1Y135-009), E, basal view (1Y135-008), F, outer-lateral view, close up showing rounded boring hole on the surface ([Y135- 010); G-H, AM F.136572, YYF4, G, inner lateral view (1Y 140-037), H, close up showing fine surface striae ([Y140-038). I-K, Pb1 element; I, AM F.136573, YYF4, outer-lateral view (1Y 135-012); J-K, AM F.136574, YYF4, J, inner-lateral view (LY 140-031), K, outer lateral view (LY 140-030). Scale bars 100 pm
unless otherwise indicated
Description
M element with a long, denticulate inner-lateral process bearing five short and widely spaced denticles (Fig. 16B), and a short, outer lateral process bearing two small rudimentary denticles; cusp robust, antero- posteriorly compressed with a sharp costa along the iner-lateral and outer lateral margins and distally curved posteriorly.
Sa element alate (Fig. 16C-F), with a robust cusp and a long denticulate lateral process on each side; cusp strongly compressed antero-posteriorly, with a sharp costa along the lateral margins; lateral process long, bearing three or more peg-like denticles (Fig. 16C), which are also strongly compressed antero- posteriorly, basal cavity open and shallow, flared posteriorly, isosceles-triangular in basal view (Fig.
Proc. Linn. Soc. N.S.W., 131, 2010
16D-E); basal margin gently arched in posterior view (Fig. 16C).
Sb element digyrate, like Sa but asymmetrical (Fig. 16G); cusp robust and antero-posteriorly compressed with sharp lateral margins; denticulate lateral process on each side bearing two or three short widely-spaced denticles; inner lateral process longer and more downwardly extending.
Sc element bipennate, strongly asymmetrical with a robust cusp, denticulate anterior and posterior processes (Fig. 16J-K); cusp distally curved inner laterally with a more convex outer lateral face bearing a prominent costa; posterior process longer and slightly arched bearing three widely-spaced denticles; anterior process curved inward bearing two widely- spaced denticles.
69
LATE ORDOVICIAN CONODONTS FROM TASMANIA
Sd element digyrate? with a robust cusp, a long denticulate lateral process on each side, a sharp costa on the posterior face and a broad anterior face with a weak carina (Fig. 16H-I); cusp with a sharp costa on each side and on the posterior face, and ornamented with fine striae; inner lateral process longer bearing eight small denticles.
Pa element blade-like with a prominent cusp, and denticulate anterior and posterior processes (Fig. 17A-H); cusp suberect or slightly inclined posteriorly, laterally compressed, standing higher above the adjacent denticles, and about twice width of the adjacent denticles on the anterior process, and typically leaving a prominent notch between cusp and the first denticle on the anterior process (Fig. 17A, G); anterior process higher and longer bearing four to eight larger and basally confluent denticles (Fig. 17A, D, G); posterior process slightly shorter, triangular in outline in lateral view, with a tapering distal end and bearing five or six smaller denticles (Fig. 17A, G); basal cavity shallow, flared laterally, forming a shallow groove underneath each process (Fig. 17E), and with a straight basal margin (Fig. 17D); some specimens bearing fine rounded boring holes (Fig. 17F).
Pb1 element asymmetrical with a suberect, robust cusp and long denticulate anterior and posterior processes (Fig. 17I-K); cusp curved inward, diamond- shaped in cross section with a sharp costa along the anterior and posterior margins, a mid costa on the inner lateral face (Fig. 17J), and a broad carina on the outer lateral face (Fig. 17K); two processes bearing small, discrete denticles; posterior process longer with six or more denticles, and anterior process shorter, extending downwards (Fig. 17K).
Pb2 element pastinate, with a robust cusp and denticulate anterior, posterior and outer lateral processes (Fig. 16A); cusp laterally compressed with a sharp costa along anterior and posterior margins and on the outer lateral face; long anterior and posterior processes bearing short, widely spaced denticles; outer lateral process short, represented by a single denticle.
Discussion
This species differs from 7. careyi in having a Pa element with shorter and higher outline bearing a prominent cusp and a notch in front of the cusp, and in having the S, Pb1 and Pb2 elements bearing small, discrete or widely-spaced denticles on the processes. Additional specimens from the Settlement Road section of Florentine Valley area confirm that it represents a separate species of Zasmanognathus. However, as only a small number of specimens are
70
available for study, this species is retained herein under open nomenclature pending further collecting and study.
ACKNOWLEDGMENTS
Field work in Tasmania by Zhen was supported by a grant from the Betty Mayne Scientific Research Fund of the Linnean Society of New South Wales. Burrett’s study was funded by the Australian Research Council. Gary Dargan (Geological Survey of New South Wales) assisted with acid leaching and residue separation. Scanning electron microscope photographs were prepared in the Electron Microscope Unit of the Australian Museum. We thank Stephen Leslie and John Pickett for their perceptive and constructive reviews of the manuscript. The study was undertaken by Zhen as part of a CAS/SAFEA International Partnership Program for Creative Research Teams, and is a contribution to IGCP Project 503: Ordovician Palaeogeography and Palaeoclimate. Percival publishes with permission of the Director of the Geological Survey of New South Wales.
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Proc. Linn. Soc. N.S.W., 131, 2010
Stratigraphic Revision of the Hatchery Creek Sequence (Early-
Middle Devonian) Near Wee Jasper, New South Wales
JAMES R. HUNT AND GAVIN C. YOUNG
Research School of Earth Sciences, The Australian National University, Canberra ACT 0200, Australia
Ghunt595@gmail.com) (Gavin. Young@anu.edu.au)
Hunt, J.R., and Young, G.C. (2010). Stratigraphic revision of the Hatchery Creek sequence (Early-Middle Devonian) near Wee Jasper, New South Wales. Proceedings of the Linnean Society of New South Wales 131, 73-92.
A new formation (the Corradigbee Formation) is erected for the upper part of the previous ‘Hatchery Creek Conglomerate’, which is elevated to Group status, its lower part renamed the Wee Jasper Formation. The “Hatchery Creek Conglomerate’, south of Burrinjuck Dam and 50 km northwest of Canberra, was previously defined as a 2.9 km thick sedimentary sequence of conglomerate, sandstone and shale nonconformable on underlying Lower Devonian limestones. The coarser lower part (Wee Jasper Formation) is now estimated at about 1500 m thick; an additional type section is nominated for its upper part, which was not included in the original type section, and lithologies, subdivision, and contacts with underlying and overlying formations are described. The upper sequence of dark shales and mudstones (Corradigbee Formation) has an estimated thickness of about 260 m, with 15 fining-upward cycles in which 50 new fossil sites have been found. Repetition of lower strata of the Hatchery Creek sequence in the west, due to an unrecognised syncline axis through the central part of the outcrop area, had suggested a much greater thickness than interpreted in this study. The relatively high topography of the softer shales and mudstones in the core of the syncline is a transient inverted topography resulting from recently eroded Tertiary basalts. The whole sequence is
interpreted as conformable on underlying limestones, and of Emsian-Eifelian age.
Manuscript received 30 October 2009, accepted for publication 17 February 2010.
KEYWORDS: Corradigbee Formation, Emsian-Eifelian, Hatchery Creek Group, Wee Jasper Formation.
INTRODUCTION
The previously named ‘Hatchery Creek Conglomerate’ is a thick sedimentary sequence of Devonian non-marine strata located 5Okm NW of Canberra (Fig. la). It is exposed over an area of about 70 km?, with most of its outcrop on the Brindabella 1:100 000 sheet, about 4 km? of which is covered by remnant Tertiary basalt (Owen and Wyborn 1979), and a small northern extension on the Yass 1:100 000 sheet (Cramsie et al. 1978). Underlying marine limestones of the Murrumbidgee Group, in the Goodradigbee valley near the village of Wee Jasper (Fig. 1b), contain an abundant invertebrate fauna, including conodonts, brachiopods, and corals (see Pedder et al. 1970, and references therein). These
provide a late Early Devonian (Emsian) maximum age limit for the Hatchery Creek sequence.
The ‘Hatchery Creek Conglomerate’ was originally assumed to be Upper Devonian in age, based on lithological similarity with the Hervey Group of central New South Wales (Pedder 1967, Conolly, in Packham 1969, Pedder et al. 1970). However a fossil fish assemblage discovered during geological mapping by Owen and Wyborn (1979) was described by Young and Gorter (1981) as probably late Eifelian (Middle Devonian) in age.
Previous authors, when referring to the ‘Hatchery Creek Conglomerate’, commented on the most accessible lower section, formed predominantly of cycles of massive conglomerate and sandstone. The measured section of Owen and
STRATIGRAPHIC REVISION OF THE HATCHERY CREEK SEQUENCE
149 v Yass
| setae. @,. Burrinjuck Dam 3 a | JX f) yy Study se A \ \
[\ Gundagai
¢ q Tumut y=
Lake George
50km
Type Section [upper Wee Jasper Fm]
__Type Section [lower Wee Jasper Fm] -(Owen & Wyborn 1981)
Inferred syncline axis \(Hood & Durney 2002) :
. Wee Jasper
[_|corradigbee Fmn ell Hatchery )
[= |Wee Jasper Fmn (WJF Creek Gp
[| Murrumbidgee Gp Limestones Burrinjuck Granite Complex
DEVONIAN Early - Middle
Peppercorn ——a “ |Beds 1km
Middle |
Figure 1. a. Regional locality map showing the study area. b. Generalised geological map showing the outcrop area of the Hatchery Creek Group, based on the Owen and Wyborn (1979) Brindabella 1:100 000 geological map, updated by detailed field mapping (e.g. eastern areas of basalt; large area to west not remapped). Previous fossil localities are the original fish locality at Windy Top (WT) described by Young and Gorter (1981), and a second fish-plant locality (JF) studied by Francis (2003). The syncline axis as identified in this study (on the left) is compared with the position of this structure inferred by Hood and Durney (2002). Boxed study areas are shown in more detail in Figs. 2-4 as indicated.
74 Proc. Linn. Soc. N.S.W., 131, 2010
J.R. HUNT AND G.C. YOUNG
Wyborn (1979) did not reach into the upper sequence above the lower massive conglomerates (Figs. 1b, 2a). The fossil fish assemblage of Young and Gorter (1981) occurs within the upper finer sequence of siltstones and mudstones, in which almost no conglomeratic horizons are seen. In this paper this upper sequence is separated out as the new Corradigbee Formation, described below, and the lower coarser sequence is renamed the Wee Jasper Formation, both formations included in the Hatchery Creek Group.
A second fossil locality (plants) was recorded on the geological map of Owen and Wyborn (1979). In 1988 an ANU student excursion located fish remains about 4 km south of the original fossil fish locality (locality 59, Fig. 3a), and apparently higher in the sequence. However the faunal composition seemed identical to that from the original fish locality, suggesting problems with the stratigraphy and structure. The plant locality of Owen and Wyborn (1979) was investigated by Francis (2003), where _ fish were found in association, this locality (JF, Figs. 1b, 2a, 3a, 4b, 5a) being only slightly higher in the sequence than the original fish locality, now called ‘Windy Top’ (WT, Fig. 1b). Hunt (2005, 2008) conducted a detailed field study of the upper fine-grained sequence (Corradigbee Formation), and discovered many additional fossil localities (Fig. 3a), mainly fish and plant remains, but with a few invertebrates (gastropods, and probable arthropods; see Appendix). New fish taxa in these assemblages (Table 1) include several osteichthyans (bony fish), and a new placoderm genus probably belonging to the arthrodires (Hunt and Young, in press; Young et al. 2010, fig. 4A). Fifteen fining-upward sedimentary cycles were identified, comprising about 260 m of the Corradigbee Formation. The cycles were mapped on both sides of the axis of a broad syncline, a major structure not shown on the geological map of Owen and Wyborn (1979). As a result their estimated total thickness of at least 2900 m for the entire sequence is erroneous. The results presented here conform closely with the first geological investigation of the area, in an unpublished honours thesis by Edgell (1949).
The original fish locality was estimated at about 1.9 km above the base of the sequence, and it was suggested that any disconformity with the underlying limestones was of short duration (Owen and Wyborn 1979; Young and Gorter 1981). Previously, Edgell (1949) had interpreted a conformable boundary between the Hatchery Creek sequence and the underlying limestones, an interpretation now followed here (see below).
Physiographically, the Hatchery Creek area of outcrop is part of the ‘Bimberi-Brindabella Upland’
Proc. Linn. Soc. N.S.W., 131, 2010
of Owen and Wyborn (1979, fig. 5), across which Miocene basalts spread into the mapped area from the ‘Kiandra Tableland’. The higher relief of the softer mudstone sequence in the “middle ridge’ of the mapped area of Hunt (2005, 2008; Fig. 3a) probably results from inverted topography. It coincides with the syncline axis, the topographic expression of which has evidently been masked by recent erosion of the cover of Tertiary basalt. Probably the basalt flowed down a previous valley representing the eroded core of the syncline, the basalt cover then inhibiting further erosion until it was eventually stripped off. A small residual cap of basalt remains adjacent to the original fossil fish locality at ‘Windy Top’ (~700 m elevation, Fig. 1b), with larger outcrops 3-5 km to the south and west (Owen and Wyborn 1979). A flagstone quarry at about 760 m elevation is located in the basalt that forms the highest part of the middle ridge of the mapped area, including Goodradigbee Hill (803 m; Fig. 3a). The area of finer sedimentary rocks was cleared for grazing many years ago, in contrast to the timbered ridges to the east in the coarser sandstone and conglomerates lower in the Hatchery Creek sequence, but since completion of this study has been revegetated as plantation pine forest.
Original access to the main outcrop was up the Cave Creek Road (locked from 2008) and along the ‘Main Ridge Trail’ to the north, then west along the “Windy Top Trail’ to the original fish locality. Access to ‘Corradigbee’ homestead (Fig. 3a) is off the access road to the 330kv power transmission line, from the south via the Tumut Road.
METHODS
Reconnaissance mapping of the lower part of the Hatchery Creek sequence by Young (1969) has been reinvestigated during many excursions to collect fossils following the research of Young and Gorter (1981), and associated with the honours project of Francis (2003). The detailed study of Hunt (2005) involved about 30 days field work on the Corradigbee Formation, covering about 20 km* in the upper section of the Hatchery Creek sequence (rectangle, Fig. 1b). The softer mudstone sequence is deeply eroded by two north-flowing tributaries of MacPhersons Swamp Creek, here termed ‘eastern creek’ and ‘western creek’, separated by the prominent ‘middle ridge’ (Fig. 3a). Erosion gullies give many good exposures of the softer sediments, and improved exposure and accessibility was a result of the 2003 bushfires in the Wee Jasper area, which burnt blackberry infestations.
15
STRATIGRAPHIC REVISION OF THE HATCHERY CREEK SEQUENCE
a b LEGEND a > ~<& endpoints of type wi z sections Bi Xsee=ssX measured section os a ped etm tracks 25 25 2 4j wee eee = faults ou J °
WT, JF [J fossil fish sites A Windy Top
w > WEE JASPER FORMATION to Pee
Murr. G limesto1
8 Windy Top Trail Type Section
po UY ISL hho
Cave Creek Rd Type Section
Proc. mn. soc. Nissw. 1B, 2010
J.R. HUNT AND G.C. YOUNG
Only some of the more significant fossil material collected from many new localities has been prepared and identified. The original description (Young and Gorter 1981) documented such forms as the placoderm Sherbonaspis hillsi (Fig. 2c), which closely resembled the ‘winged fish’ first described by Hugh Miller (1841) from classic Middle Devonian Old Red Sandstone fish faunas of Scotland. This was the first discovery of such an assemblage from the Southern Hemisphere. An updated faunal list for the Hatchery Creek fish assemblage is given in Table 1; formal fossil descriptions will be presented elsewhere.
For the Corradigbee Formation, various field sites were examined as to the bedding type, dip, strike, lithology and sedimentary structures (see Appendix). Many fining-upward sedimentary cycles could be seen on air photographs by their more resistant basal sandstones, and were traced out on a 90x90 cm photo enlargement. Some identified beds were walked along strike to establish correlations between _ different exposures for the detailed stratigraphy (Figs. 2a, 4a). Sedimentary strata with good exposure were selected for measured stratigraphic sections using either a tape or 150 cm Jacobs staff and abney level. The cycle containing the original 1981 fossil fish locality (WT) was called Cycle A, with overlying cycles labelled up through the sequence as B, C, etc., and underlying cycles down the sequence labelled B’- F’. The thickness of the Wee Jasper Formation was estimated using aerial photographs and data plotted from the lowest beds of the Corradibee Formation and measured off the maps and photos.
Numbered localities are shown in Fig. 3aand listed in the Appendix. For different field investigations the locality numbers are: 1-24, 59-159 (Hunt 2005); 160- 161, 062-082 (Hunt 2008); prefix GY (Young 1969); prefix JF (Francis 2003). All grid references refer to the Wee Jasper 1:25 000 topographic map 8627-4N (second edition, 2003). Full grid references (as in appendix) are abbreviated in the text (e.g. 646385 611805 shortened to GR46385 1805). Fossil material is registered in the ANU palaeontological collection, Canberra (Building 47, Research School of Earth Sciences).
PREVIOUS STRATIGRAPHY
The ‘Hatchery Creek Conglomerate’, named by Joplin etal. (1953), consists of cyclothems of terrestrial conglomerates, sandstones and mudstones. These fine upwards and the beds are laterally extensive, some being traceable over several kilometres along the length of the outcrop (Young 1969). These beds can be classified as red beds according to the definition of Van Houten (1973).
Owen and Wyborn’s (1979) estimated thickness of about 2.9 km for the Hatchery Creek Conglomerate was followed by other authors (Young and Gorter 1981; Branagan and Packham 2000; Packham 2003). With the subdivision of this sequence into two formations as proposed here (the Wee Jasper Formation and the Corradigbee Formation), and the recognition that the previously interpreted upper ~300 m of coarse sandstones and conglomerates is in fact a repetition of the lower strata (Wee Jasper Formation) on the western limb of a syncline, a significantly reduced total thickness estimate of 1760 m for the Hatchery Creek Group is based on the following: thickness for the lower formation (Wee Jasper Formation) estimated from air photos (average dip 40°) at about 1500 m; thickness for the upper Corradigbee Formation (as defined below) estimated at 260 m.
HATCHERY CREEK GROUP (UPGRADED FROM FORMATION)
WEE JASPER FORMATION (NEW NAME)
The first published description (as “Hatchery Creek Conglomerate’) recorded numerous fining- upward conglomeratic cycles (Owen and Wyborn 1979: microfiche M314-M320). A type section comprising about 1200 m of almost continuous exposure of cycles of ‘conglomerate, sandstone and siltstone typical of the lower part of the formation’ was nominated along the Cave Creek Road (see Fig. 1b), from the basal contact with the underlying carbonates at their stated grid reference (GRS509 176), to the top at the T-junction of the Cave Creek
Figure 2 (LEFT). a. Detailed geological map of the Wee Jasper Formation (previously Hatchery Creek Conglomerate, lower part) between the original type section (Cave Creek Road) for the lower part de- fined by Owen and Wyborn (1979), and the new type section for the upper part (Windy Top Trail) de- scribed in the text. Coarser basal part of each fining-upward unit indicated by stippling or shading. b. Summary section for the lower 1600 m of the Hatchery Creek Group, showing correspondence between the upper cycles of the Wee Jasper Formation and lower cycles of the Corradigbee Formation. c. Recon- struction of the placoderm fish Sherbonaspis hillsi Young and Gorter (1981), which established a prob-
able Eifelian age for the Hatchery Creek sequence.
Proc. Linn. Soc. N.S.W., 131, 2010
VW
STRATIGRAPHIC REVISION OF THE HATCHERY CREEK SEQUENCE
Road and Main Ridge Trail (their GR491 172; Fig. 2a). Owen and Wyborn (1979) noted a change at about 1500 m above the base of the formation to a lithology dominated by fine buff sandstone and red siltstone with root casts. They considered but did not follow the stratigraphic subdivision first proposed by Edgell (1949), who separated off this finer upper sequence as the ‘Middle Ridge Shales’ from the lower ‘Wee Jasper Creek Conglomerates’ (also overlooked by Packham 1969; Pedder et al. 1970).
Young (1969) had previously subdivided the lower 1550 m of the Hatchery Creek Conglomerate into four units, the lower Units | and 2 forming the eastern slope of the main ridge along the western margin of the Goodradigbee valley, and the upper Units 3 and 4 mainly outcropping in the western drainage of Macphersons Swamp Creek. The top of the formation was left undifferentiated. This subdivision has been checked in the field since 2003, supported by air photo interpretation using new colour air photos, and more recently Google Earth images, as summarised in Figure 2a. Estimated thickness from the base for these four units was 250, 200, 400 and 700 m (Young 1969). Owen and Wyborn (1979) stated that the cycles as defined by the beds of conglomerate rarely extend beyond about | km, but some of the units mapped by Young (1969), for example the prominent basal conglomerates of Units | and 2, can be traced on air photos nearly 10 km along the western escarpment of the Goodradigbee valley (Fig. 2a). The basal conglomerates of Unit 2 form a row of conspicuous outcrops about one third of the distance up the slope of each spur between about GR495 210 and GR492 220. Both horizons can be traced north (with two slight fault displacements at about GR495 222 and GR492 232) at least to GR490 245. Unit 3 crops out near the top and over the ridge to the west.
To the south, prominent outcrops of three ridges north of the road in the Cave Creek Road type section of Owen and Wyborn (1979) can be assigned to the basal coarse beds of Units 1-3 (between GR509 174 and 504 171). The basal conglomerate of Unit 3 can be readily traced on air photos from GY52 (GR499 193) to a prominent knoll on the spur at GR497 197, and then to the crest of the main ridge between GR492 208 and 489 219. Farther north a sharp bend to the west in the track crosses the basal conglomerate of Unit 4 at GR4855 221. This basal conglomerate is readily traced along strike to the south as a series of prominent outcrops between valleys (e.g. GR487 2125, 487 208), and forms the first outcrop of conglomerate encountered after the turnoff into the eastern end of the Windy Top track, at GR489 2015.
Since the existing type section finishes well below
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the lithological change to much finer sediments (the base of our new formation), we nominate an additional type section for the upper part of the renamed Wee Jasper Formation, along the Windy Top Trail from its junction with the main track at GR491 201, to the vicinity of the locked gate at Windy Top (GR477 2016), about 1.4 km to the west. This is accessible by 4-wheel drive vehicle, and the valleys to the north and south display a thick section of alternating coarse and fine beds as mapped by Young (1969). From the eastern end of this type section, down the spurs into the Goodradigbee valley, air photos clearly show the base of Unit 3 at GR494 201, the base of Unit 2 at GR496 2065, and the base of the Hatchery Creek Group (and Unit | of the Wee Jasper Formation) on the edge of the treeline at GR5012 202.
Owen and Wyborn (1979) recorded a fine- grained sequence between about 1500-2600 m above the base of their Hatchery Creek Conglomerate, then a return to cyclic conglomerates about 300 m thick at the top of the sequence. However our more detailed mapping has shown this interpretation to be incorrect, these ‘upper’ conglomerate cycles in fact representing a repetition of the contact between the Wee Jasper Formation and the Corradigbee Formation on the western limb of the syncline. The western contact (running beneath the largest basalt outcrop; Fig. 1b) was not mapped in detail, but approximates to the corresponding formation boundary of Edgell (1949). The most westerly discovered fossil site (Fig. 3a, locality 160; with fish and plants) is still in the Corradigbee Formation. Further west, light yellow sandstones of the Wee Jasper Formation were observed in the vicinity of GR449 174, but to the north similar horizons are more conglomeratic where they emerge from beneath the basalt (near GR450 203). A similar increase in coarseness to the north was observed on the eastern limb of the syncline (see below). The uppermost coarse layers of the Wee Jasper Formation are exposed within the main outcrop of the Corradigbee Formation, in the creek bed along a section of the Western Creek (dashed line, Fig. 5a), but too narrow to be shown on the geological map (Fig. 1b). Here, the lower levels of the Corradigbee Formation beneath measured section 2 (see Fig. 3) are inaccessible with a steep drop down to the creek bed.
Lower and Upper Contacts
Various authors have commented on the nature and significance of the contact between the Hatchery Creek sequence and the underlying marine limestones, but only some of these were based on actual field © investigations. Young (1969, p. 47) discussed the
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J.R. HUNT AND G.C. YOUNG
upper limestone boundary, noting that the uppermost Unit 6 of his ‘Upper Reef Formation’ was generally poorly exposed because of high clay content, and was covered by scree from the much more prominent overlying “Hatchery Creek Conglomerate’ (now Wee Jasper Formation). Where Unit 6 had continuous exposure on the western shore of Lake Burrinjuck, north from about GR491 243 around to the mouth of Hatchery Creek, the beds were highly sheared in the vicinity of the fold axis. The same applies at the southern fold closure in the vicinity of the Long Plain Fault south of Wee Jasper, obscuring sedimentary changes at the boundary.
Young (1969) noted there was no change of strike across the boundary, and no limestone clasts were observed in the basal conglomerate. However, in four measured sections across this interval there was a marked difference in thickness of the uppermost Unit 6, from 80 m in the south at GY39 (GR520 136), 210 m at GY40 (GRS508 183), 140 m
_at GY43 (GR499 210), and 110 m at GY44 (GR494
230). This thickness variation was attributed to slight warping (less than 1°) before deposition of the conglomerate, indicating a disconformable contact. Pedder et al. (1970, p. 210) independently provided similar evidence for a disconformable contact, noting that the “Hatchery Creek Conglomerate’ (Wee Jasper Formation) on the eastern limb ‘rests more than 250 feet above the highest assemblage zone of the Taemas Formation, whereas on the western limb it may rest less than 100 feet above the Hexagonaria smithi smithi Teilzone’. They also noted that ‘the lithologies of the two formations belong to entirely distinct megafacies’. Owen and Wyborn (1979, M320) also favoured a disconformable contact on the evidence of thickness variation in the uppermost unit of the Taemas Limestone, but suggested, from the age evidence of the overlying fish assemblage (subsequently published by Young and Gorter 1981), that a ‘disconformity — if present — represents a short time duration’.
Subsequent to these field investigations a new track was cut around the western shore of the lake at the northern end of the Goodradigbee valley. This gave much improved exposure of this contact in the vicinity of GR488 252, an important fossil fish locality in the limestone (Fig. 2a). Here, Campbell and Barwick (1999) measured a section through the contact, the uppermost beds of the Taemas Limestone comprising about 110 m of thin-bedded limestones and shales ‘interpreted as an intertidal zone carbonate deposit consistent with the fact that the overlying unit is the fresh water Hatchery Creek Formation’ (p. 125). Lindley (2002, fig. 4) presented a revised version
Proc. Linn. Soc. N.S.W., 131, 2010
of this section, with the uppermost unit beneath the conglomerate assigned to Unit 6 of the ‘Upper Reef Formation’ of Young (1969), and Campbell et al. (2009, p. 62) noted that the top of carbonate sequence with shallow marine algal mats was ‘transitional into the overlying fresh water Hatchery Creek Formation’.
Although uncertainty about this boundary was indicated in stratigraphic sections of Basden et al. (2000, fig. 2) and Young and Turner (2000, fig. 3B), the new evidence just summarised is accepted as indicating a conformable contact at the base of the Hatchery Creek Group. The thickness variations in the uppermost limestone units noted above must therefore be interpreted as a depositional feature. This complies with the original opinion of Edgell (1949, p. 10) that interbedded lithologies at the contact indicated continuous deposition.
The upper contact of the Wee Jasper Formation (and base of the new Corradigbee Formation as defined below) is at the top of Cycle D’ of Hunt (2005). This is the highest cycle observed with conglomerate/coarse pebbly sandstone forming the basal unit, all higher cycles having sandstone at the base (the rare thin conglomerates described below for the Corradigbee Formation were within a cycle, not at the base). It is noted that coarse beds persist to the top of the Wee Jasper Formation in the vicinity of localities 062 and 068 (Fig. 2a), but farther south the equivalent beds seem less coarse, the contact being less clearly defined, and recognised by a change in colour rather than grainsize (discussed below).
Subdivision
The general outcrop of the Wee Jasper Formation is indicated in Figure 1b, and a refined version of Young’s (1969) subdivision into four units is detailed in Figure 2. As noted above, the coarser basal unit of each cycle (normally about 30-40 m thick), can generally be traced with confidence on air photos, although individual beds may pinch out along strike. For example a prominent ridge just west of the Main Ridge Trail at GR495 190 (Fig. 2a) is the next resistant set of beds above the base of Unit 3, it forms the main ridge for about 1 km along the track to the south, but is less clearly differentiated in the Cave Creek type section (Unit 3a, Fig. 2a). To the north it is traceable to a similar prominent ridge immediately east of the track at GR492 199, and it also crosses the track at the Windy Top Trail turnoff. It forms prominent outcrops immediately west of the track between GR490 208 and 4895 213, before it is crossed by the track again at about GR488 219, where it is less distinct. This is a distance of about 3 km along strike for what
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STRATIGRAPHIC REVISION OF THE HATCHERY CREEK SEQUENCE
Table 1. Faunal list for the Hatchery Creek fish assemblage (updated from Young and Gorter 1981).
Agnatha Thelodontida
1. Turinia sp. cf. T. hutkensis Blieck & Goujet (Young & Gorter 1981)
Gnathostomata
Acanthodii
2. climatiid gen. et sp. indet. ?diplacanthiform gen. et sp. indet.
3 4. Tareyacanthus sp. cf. T. magnificus Valiukevicius (Burrow 2002) 5
Watsonacanthus? sp.
Osteichthyes (Sarcopterygii)
6. Gyroptychius? [new genus] australis Young & Gorter, 1981
7. osteolepiform gen. et. sp. nov. 2 (Hunt 2008) 8. osteolepiform gen. et. sp. nov. 3 (Hunt 2008) 2.
?onychodontid indet.
Placodermi Arthrodira
10. Denisonosteus weejasperensis Young & Gorter, 1981
11. cf. Denisonosteus sp. nov. (Hunt 2005)
12. coccosteomorph cf. Coccosteus (Hunt 2008)
13. ?arthrodire gen. et. sp. nov. Hunt and Young, in press.
14. Arthrodira incertae sedis
Antiarcha
15. Sherbonaspis hillsi Young & Gorter, 1981 16. cf. Sherbonaspis sp. nov. (Hunt 2005)
17. Monarolepis verrucosa (Young & Gorter 1981) Young, 1988
is interpreted as a laterally discontinuous coarser interval in the middle part of Unit 3.
The overlying recessive zone, representing the top of Unit 3 at its boundary with the basal conglomerate of Unit 4, is more persistent along strike, being traceable over about 5 km back to the Cave Creek Road type section. In the north it is crossed at a sharp turn in the Main Ridge Trail at GR4855 221, it can be followed south to GR4893 2015 (Windy Top Trail), GR490 1955 (next valley south), GR4955 180 (east- west section of Main Ridge Trail), and GR4955 1705 (Cave Creek Road type section).
Above this in the Cave Creek Road type section, the coarse basal part for the overlying Unit 4 as mapped by Young (1969) corresponds to a sharp bend in the Cave Creek road at GR495 170. Unit 4 is subdivided into 9 fining upward cycles (4a-j), the upper parts of which correspond to the five ‘thin zones of low weathering resistance’ mapped by Young (1969). These are readily identified on recent
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air photos in the valleys to the north and south of the Windy Top Trail, designated here as type section for the upper part of the Wee Jasper Formation. The basal conglomerate/pebbly sandstone of Unit 4 (cycle 4a) is about 40-50 m thick, fining up into a poorly outcropping interval of similar thickness, the latter clearly visible on air photos as a continuous less resistant zone from GR4845 224 south to the Windy Top Trail type section. Here it separates the basal conglomerate of Unit 4 at GR489 2015, and the basal coarse beds of the second cycle, encountered at the first bend in the track (GR488 202). This is the lowest of three similar fining upward cycles (4b-d) crossed by the track before a sharp southerly bend at GR4935 202. Each cycle is estimated at about 70 m thick, with the coarse resistant beds comprising more than half the thickness (4b, c), or about half (4d). These three units are well exposed in the next creek to the south, between about GR485194 and 490 196.
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On air photos (and ‘Google Earth’) the E- W sections along the valleys of the three creeks to the north of the Windy Top Trail clearly show the alternating resistant and five recessive beds of Unit 4 as mapped by Young (1969). The undifferentiated upper part of the ‘Hatchery Creek Conglomerate’ of Young (1969) approximates to the Corradigbee Formation as defined below. The upper part of cycle Ac is the lowest of the five ‘less resistant mudstones’ mapped by Young (1969), and can be traced to the north at least as far as the vicinity of GR478 222. The recessive upper part of cycle 4d thickens along strike to the north of the Windy Top Trail, in the vicinity of GR483 205. The overlying four cycles (4e-h) in this valley (the first creek north of the track, between GR490 205 and GR475 206) are seen as narrow ridges separated by less resistant bands of equal or greater width. Most can be traced farther north to the valley section of the creek between GR476 216 and GR487 216, where the resistant bands are thinner _and recessive bands correspondingly thicker. The base of cycle 4e is traceable to the south to cross the Windy Top Trail immediately west of the sharp bend at GR483 200. Where the northern creek turns to the north-west at GR476 216 the creek has eroded along the upper recessive bed mapped by Young (1969). This is the upper part of cycle 4f, traceable back to GR481 2005 on the Windy Top Trail. The basal coarse bed of cycle 4g is the lowest of three apparently thicker fining- upward cycles (4g,h,j) along the Windy Top Trail, their finer upper parts forming gullies immediately to the south. However further south between about GR475 194 and GR482 194 these beds are more differentiated, and the less weathered outcrop along the track may be due to relatively recent exposure by removal of the overlying basalt. The uppermost of these units (4j) passes beneath the remnant basalt cap of Windy Top (Fig. 4b).
The correspondence between the uppermost