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Late Triassic megaspores from the AmeryGroup, Prince Charles Mountains, EastAntarcticaDAVID J. CANTRILL a & ANDREW N. DRINNAN aa School of Botany, The University of Melbourne , Parkville, Victoria,3052, AustraliaPublished online: 23 Sep 2010.

To cite this article: DAVID J. CANTRILL & ANDREW N. DRINNAN (1994) Late Triassic megaspores fromthe Amery Group, Prince Charles Mountains, East Antarctica, Alcheringa: An Australasian Journal ofPalaeontology, 18:1-2, 71-78, DOI: 10.1080/03115518.1994.9638765

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LateTriassic megaspores from the Amery Group,Prince Charles Mountains, East Antarctica

DAVID J. CANTRILL AND ANDREW N. DRINNAN

CANTRILL. D. J.. & DRINNAN. A. N.• 1994:03:28. Late Triassic megaspores from the AmeryGroup. Prince Charles Mountains. East Antarctica. Alcheringa 18.71-78. ISSN 0311-5518.Megaspores referable to the genera CabochonicusBatten & Ferguson 1987 and Minerisporites

Potonie 1956 are a common component of the palaeoflora recovered from the Jetty Member withinthe Flagstone Bench Formation of the Amery Group. The known ranges of these two genera. inconjunction with the macro floral remains. suggest a Late Triassic age. Two new species.Cabochonicussinuosus and Minerisporitestriangulatus, are described. Scanning electron micro­scopic examination of the spores indicates that standard palynological treatment of megasporescan result in sculptural degradation. possibly leading to incorrect generic assignment.

D. J. Cantrill and A. N. Drinnan 2, School of Botany, The University o[ Melbourne, Parkville,Victoria 3052, Australia. (lpresent address: British Antarctic Survey, Natural EnvironmentResearch Council, MadingleyRd, High Cross, Cambridge, CB30EI; United Kingdom. 2Author[or correspondence); received 8 December 1992.

Keywords: Megaspores. Cabochonicus, Minerisporites , Late Triassic. Antarctica.

PALAEOZOIC sediments in East Antarcticaoutside the Transantarctic Mountains were firstrecognized in the northern Prince CharlesMountains around Beaver Lake, where flatlying, arkosic sandstones crop out (Crohn,1959). Crohn (1959) named the sequence theAmery Formation and tentatively assigned anUpper Permian age. Subsequent palynologicalexamination of material collected on variousAustralian National Antarctic Research Expe­ditions yielded poorly preserved palynomorphsthat confirmed a Late Permian age (Balme &Playford, 1967; Kemp, 1973; Playford, 1990).Macroremains of Glossopteris Brongniart, ~r­tebraria Royle and silicified wood of theAraucarioxylon type also supported the Permiandetermination (White, 1973), Mond (1972)raised the status of the Amery Formation togroup rank and recognized three formationswithin the sequence; Radok Conglomerate,Bainmedart Coal Measures and FlagstoneBenchFormation. A Triassic age was suspectedfor the upper part of the sequence (Ravich et

031115518/94/010071-08 $3.00 ©AAP

al., 1977), but this was not confirmed untilrecently when further field work revealed thepresence of the corystosperm foliage Dicroid­iumzuberi (Szajnocha) Archangelsky within theFlagstone Bench Formation on Jetty Peninsula(Webb & Fielding, 1993). Field work in the1991/92 season to collect Late Permian petri­fied peat and additional Triassic material re­sulted in the recognition of a number of newlocalities on Jetty Peninsula (Fig. 1). Bulkmaceration of this material for cuticular frag­ments revealed an abundance of megaspores,which form the basis of this paper.

Examination of spores and pollen by lightmicroscopy relies on the palynomorphs beingtranslucent. To achieve translucence, spore res­idues are macerated by an oxidative step fol­lowed by removal of the oxidation products withalkali or bleach (Traverse, 1988), This processhas long been accepted by palynologists despitethe fact that spores were often observed toshrink or expand during this process. Theadvent of electron microscopy allowed the ex­amination of spores in relatively unmodifiedstates after extraction from the rock matrix.

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72 D.1. CANTRILL & A. N. DRINNAN ALCHERINGA

KAMENISTAYArJ-70030'PLATFORM '1! .

~

-70045'

I690

10km

ELSEPLATFORM.",.",_,,,

f)

\.@ ~,~" ...

BEAVER LAKE i

@~ .....,.....i'

Isao

[I Flagstone Bench Formation

D Bainmedart Coal Measures

~ Radok Conglomerate

~ Precambrian

LambertGlacier

NORTHERNPRINCECHARLESMOUNTAINS

.; !,'.

Fig. 1. Beaver Lake area of the northern Prince Charles Mountains. east Antarctica. Spore symbol = Triassic megasporelocalities.

Materials and methodsThe sediments consist of light to dark grey,planar laminated, micaceous sandstones andsiltstones that contain abundant fragments ofDicroidium zuberi and conifer shoots of theform-genus Pagiophyllum, and rare fragmentsof D. crassinervum var. stelznerianum (Geinitz)Anderson & Anderson, Pteruchus dubiusThomas emend. Townrow, conifer cones, andcycad-like foliage (Webb & Fielding, 1993;Cantrill, Drinnan & Webb, in prep.). The rockwas bulk macerated in hydrofluoric acid,washed in distilled water and passed through a100 um mesh sieve. Some megaspores werepicked out during examination and mounteddirectly onto SEM stubs. Stubs were dehy­drated overnight and coated with gold/palla­dium for examination in a lEaL lSM-840 SEM.

Other megaspores were prepared for micros­copy using standard palynological techniques oftreatment in either nitric acid or Schulze's solu­tion, and/or dilute ammonia. The effect of thedifferent treatments was compared using SEM.

Megaspores for TEM were dehydrated in agraded series of acetone before being embeddedin Spurr's resin (Spurr, 1969). They werestained for 7 minutes in saturated uranyl acetatein water and 1 minute lead citrate, or for 10minutes in saturated uranyl acetate in 30%ethanol and 1 minute lead citrate. The sporeswere examined in a lEaL 1200EX TEM at80KV.

Megaspores are mounted on three SEM stubsdeposited in the palaeontology collection of theNational Museum of Victoria (NMV). Onlyillustrated specimens have been allocated NMVnumbers.

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ALCHERINGA ANTARCTIC TRIASSIC MEGASPORES 73

Cabochonicus Batten & Ferguson 1987

Cabochonicus sinuosus sp. nov. (Fig. 2)

Systematic palynology

Holotype. NMVP197989 (Fig. 2A).

Other specimens. NMVP197990,NMVP197991, ten unnumbered specimens.

Remarks. Cabochonicus was erected for spe­cies previously referred to Verrutriletes van derHammen on the basis that the reddish coloured,resin-like, gemmate ornamentations were nottrue verrucae but represented exinal excres­cences (Batten & Ferguson, 1987). This sporeornamentation had also previously been inter­preted as fungal in origin (Marcinkiewicz,1979). Banerji et al. (1984) supported thefungal interpretation, as the ornamentation dis­solved in acid and so was considered to have adifferent composition to the exine. In contrastWaksmundzka (1985) examined gemmate struc­tures on Verrutriletes imitatus Dijkstra 1961 andV. guttatus Marcinikiewicz 1971 by TEM andconcluded that they were part of the sporeornamentation, as sporopollenin threads werecontinuous with the verrucae. This was furthersupported by Batten & Ferguson (1987) whoexamined Verrutriletes (now Cabochonicustcarbunculus Dijkstra 1949 and interpreted theornamentation as exinal outgrowths. Examina­tion of thin sections using transmission electronmicroscopy (Fig. 2F) shows these structures arecomposed of sporopollenin, and are continuouswith the outermost sporoderm layer. Interpre­tation of this outer sporoderm layer is problem­atical, as it is uncharacteristic of extantpteridophyte megaspores. Pteridophyteisospores have a similar, thin, outer layer ofamorphous sporopollenin that differs structur­ally from the rest of the exospore and is de­stroyed by harsh acetolysis. Perispore layers,which are also absent in extant pteridophytemegaspores, have the same hardy resistance toacetolysis as the inner exospore (Tryon &Lugardon, 1990). The destruction of the outersporoderm layer and ornamentation inC. sinuosus by acid treatment suggests it is atrue exospore layer with attached, or closelyadhering, tapetum-derived globules.

The dissolution of Verrutriletes ornamenta­tion in acid led Banerji et al. (1984) to note thesimilarity of the oxidized spores to Banksispor­ites kachchhensis Banerji, Jana & Maheshwari1984. Similarly, Batten & Ferguson (1987)

verrucae are sparse (Baldoni & Taylor, 1985),C. guttatus has verrucae that are highly variablein size (Waksmundzka, 1985; Batten & Fergu­son, 1987). In contrast, C. sinuosus has verru­cae on all surfaces, including the contact faces.

Verrutn'letes carbunculusType species.Dijkstra 1949.

Diagnosis. Trilete megaspore, amb circular tosubcircular, ranging from 150 to 300 fiITl indiameter. Laesurae extending from 0.5 to 0.7of the spore radius, sinuous, lipped (Fig. 2A).Lips laevigate, 8 to 20 !-lm broad and up to 10fJ.m high at pole but tapering to 2 !-lm. Outer­most sporoderm layer up to 2 !-lm thick,micropitted, surface appearing finely reticulate(Fig. 2C). Ornamentation comprising sparse,gemmate or rarely clavate, lustrous, reddish­brown structures, 2 to 10 !-lm in diameter, 1 to10 fiITl tall, on the equatorial and distal surfaces(Figs 2B, C, F), and occasionally on the proxi­mal contact faces (Fig. 2A). Occasional sporesalmost devoid of sculpture. Middle sporodermlayer up to 10!-lm thick, composed of reticulate,labyrinthine network of sporopollenin threads(Fig. 2D). Inner sporoderm layer thin, lami­nated, often shrunken into the centre of thespore (Fig. 2G).

Comparison. Batten & Ferguson (1987) as­cribed five species to Cabochonicus: C. im­itatus (Dijkstra) Batten & Ferguson 1987,C. pseudosquamosus (Marcinkiewicz) Batten& Ferguson 1987, C. guttatus (Marcinkiewicz)Batten & Ferguson 1987, C. gamerroi (Baldoni& Taylor) Batten & Ferguson 1987 and C. car­bunculus (Dijkstra) Batten & Ferguson 1987.The most striking feature distinguishingC. sinuosus from other species of Cabochoni­cus is its sinuous laesurae. All species havedifferences in the distribution and size of theverrucae; C. imitatus and C. carbunculus lackverrucae on the contact faces but otherwise tendto have abundant verrucae (Banerji et al., 1984;Batten & Ferguson, 1987), C. gamerroi alsolacks verrucae on the contact faces but the

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noted that the type species Cabochonicus car­bunculus was mixed with laevigate megasporesreferred to Triletes murrayi (Harris)Marcinkiewicz 1971. Occasional specimens ofCabochonicus sinuosus have only one or twoglobules and closely resemble Banksisporitessinuosus Dettmann 1961 (now Triletes sinuosus(Dettmann) Fuglewicz 1973), and rare speci­mens that completely lack ornamentation matchthe description of B. sinuosus.

Dettmann (1961) recovered Banksisporitessinuosus from Rhaetian sediments from Tasma­nia by maceration in hydrofluoric acid followedby2 to 3 hours in Schulze's solution and clearingin 5% potassium hydroxide before extraction ofmegaspores. Spores of Cabochonicus sinuosuswere subjected to maceration in 70% nitric acidfor 2 to 3 hours, or Schulze's solution for 30minutes and cleared in dilute ammonia. Theresult of this treatment was the destruction ofthe outermost sporoderm layer along with theornamentation (Fig. 2E). The spores that wererecovered by this standard palynological tech­nique would be classified as Banksisporitessinuosus. Cabochonicus sinuosus andBanksisporites sinuosus are probably conspe­cific, but new preparation of material from theTasmanian localities is needed to determine thisconclusively.

Minerisporites Potonie 1956

Type species. Selaginellites mirabilis Miner1935.

Minerisporites triangulatus sp. nov. (Fig. 3)

Holotype. NMVP197993 (Fig. 3B)

Other specimens. NMVP197992,NMVP197994, ten unnumbered specimens.

ANTARCTIC TRIASSIC MEGASPORES 75

Diagnosis. Trilete megaspore, biconvex, ambtriangular to subtriangular, equatorial diameterranging from 350 to 400 IJ.m. Laesurae extend­ing slightly beyond the equator of the spore(Figs 3A, B, D). Termination of radii rangingfrom Y-shaped (Fig. 3A) to rounded (Fig. 3B),auriculate, auricles projecting beyond the zona.Zona narrow, up to 50 IJ.m high at intersectionswith laesurae, but projecting only slightly abovethe spore wall in the interradial areas. Radiistraight, lips thin, raised to form a prominentflange 30 to 60 IJ.m high, flange slightly serrate.Exospore surface with positive reticulated muri,muri up to 4IJ.m high, evenly distributed on allfaces including the zona. Exospore composedof horizontally aligned, reticulate network ofsporopollenin sheets (Fig. 3C) that are closelypacked and appearing laminated toward theinside, but are less dense toward outside.

Comparison. The spores described above areclearly referable to Minerisporites as they havea distinct zona, lack cushions along the laesuraeand have reticulated sculptural elements. Over35 species have been described from Triassic toEocene sediments (Batten & Kovach, 1990).However, the majority of species occur in Mid­dle to Late Cretaceous sediments. Only onespecies, M. ales Jung 1960, is known from theTriassic, and although a number of species aresuggested to be Jurassic/Cretaceous in age onlyM. richardsonii (Murray) Potonie 1956 emend.Harris 1961 occurs in Early as well as LateJurassic sediments (Kovach & Batten, 1989).Minerisporites ales clearly differs fromM. triangulatus; in particular it has large auri­cles ranging from 1;2 to V3 times the sporediameter and the proximal faces of the sporehave long spines, features that are unknown inM. triangulatus. Minerisporites richardsoniihas small auricles like M. triangulatus but hasabundant spines on the proximal faces. Creta­ceous species of Minerisporites occasionally

Fig. 2. Cabochonicus sinuosus sp. nov. A, proximal surface; note scattered globules and slightly sinuous lacsurae,NMVP197989 (Holotype). B, distal surface with scattered globules, NMVP197990. C, surface view illustrating thin outersporodcrm layer with globules. and middle sporoderm layer. D. fractured surface of spore wall in transverse sectionillustrating the loosely attached outer sporodcrm layer; exospore composed of reticulate sporopollcnin network. E. sporeafter treatment in nitric acid: note complete destruction of outer sporoderm layer and ornamentation, NMVP197991. F,TEM of section through globule showing continuity with outer sporoderm layer. G. TEM of section through innermostexospore layer. which retains a compressed, but foliated. substructure. Scale Bars A, B, E = 100!J1Tl. C. D, F = 10 um,G = 1!J1Tl.

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Fig. 3. Minerisporites triangulatus sp. nov. A. B. proximal view. A. NMVP197992. B, NMVPI97993 (Holotype), C,TEM of section through muri with degradation of spore wall at apex of muri. Wall is composed of a network of anastomosingfoliated sheets that are compacted towards the inside of the spore wall. D, equatorial view, NMVP197994. Scale bars A.B, D = 100 fJm. C = 10 px«.

appear similar to M. triangulatus, however, thereticulated muri are generally coarser (e.g.M. mineri (Sukh-Dev) Banerji, Jana &Maheshwari 1984, M. reticulatus (Singh,Srivastava & Roy) Banerji, Jana & Maheshwari1984), the termination of the auricle is notY-shaped (M. auriculatus Singh, Srivastava &Roy 1964), or the laesurae may be sinuous(M. cutchensis Singh, Srivastava & Roy 1964).Thus, although the specimens described here

can be assigned to Minerisporites they do notconform to any of the existing species, and sohave been assigned to a new species,M. triangulatus.

Discussion

Affinities. Production of megaspores that areshed from the parent plant is today confined to

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ligulate -lycophytes and water ferns(Marsileales, Salviniales). Megaspores refera­bleto Banksisporites Dettmann 1961 have beenrecovered from lycopodiaceous cones (e.g,Helby & Martin, 1965) and the similarity ofCabochonicus sinuosus to Banksisporites sug­gests that these spores are also lycopodiaceous.Batten & Ferguson (1987) tentatively suggesteda greater affinity of Cabochonicus toSelaginellales based on the external morphol­ogy of the spore and the wall stratification,howeverthey did not exclude the possibility thatthey were derived from a pleuromeid type oflycophyte. Minerisporites type megasporeshave also been associated with lycophytes(Krassilov, 1982), and in particular Isoetales(Hickey, 1977). Isoetites horridus (Dawson)Brown from the Paleocene of North Americawas found to contain in situ Minerisporitesspores (Hickey, 1977; Melchior, 1977).

Age. The megaspores are unable to give aprecise age range for the sequence but supporta Late Triassic age, probably Rhaetian. Thegenus Cabochonicus ranges from late Rhaetianto Santonian (Batten & Ferguson, 1987),Minerisporites ranges from Rhaetian to Eocene.The similarity of acid-treated Cabochonicussinuosus to Banksisporites sinuosus, a sporerecorded from Rhaetian sediments of Tasmania(Dettmann, 1961) and Carnian-Norian strata ofthe Tiki Formation in India (Banerji et al.,1978), offers further support for a late TriassicAge. However, material from Tasmania needsto be examined to determine if Cabochonicussinuosus is in fact identical to Banksisporitessinuosus.

Macrofloral remains of Dicroidium Gothanindicatea minimum age of latest Triassic for theJetty Member sediments. Dicroidium zuberiranges from Early to Late Triassic and Dicroid­ium crassinervum var. stelznerianum rangesfrom Middle to Late Triassic age (Anderson &Anderson, 1983). Indirect evidence fromstratigraphic succession also supports a Middleto Late Triassic age. In eastern Australia(Bowen, Sydney, and Tasmanian Basins) theLate Permian to Early Triassic succession ismarked by a transition from coal-producing tored bed sedimentation. The presence of redbedsbelow the fossil locality suggests the local­ity is younger than Early Triassic. It therefore

ANTARCTIC TRIASSIC MEGASPORES 77

seems most likely, based on the palynology,macroflora and stratigraphic succession, thatthis part of the sequence is Late Triassic in age.

AcknowledgmentsWe would like to thank Dr John Webb whoinitially aroused our interest in this project aftershowing us material he collected in the 1989/90ANARE field season. Our appreciation goes tothe Australian Antarctic Division for the logisticsupport, and in particular to Louise Crossleyand the other members of the ANARE 1991/92Prince Charles Mountains expedition. Thanksalso to Jocelyn Carpenter for preparing speci­mens for TEM. Anonymous reviewers pro­vided helpful comments on the manuscript.Laboratory expenses were funded by grantsfrom the Antarctic Science Advisory Commit­tee and Australian Research Council.

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