Tertiary occurrence of the fern Lygodium (Schizaeaceae) in Australia and New Zealand

20 Pages·1992·8.6 MB·English

by  A C Rozefelds

TERTIARYOCCURRENCEOFTHE FERNLYGODWM(SCHEAEACEAE
INAUSTRALIA AND NEW ZEALAND \
A.C-ROZEFELDS.DC CHRISTOPHELANDN.F. ALLEY
&
Kozctclds.A.C.,Christophel,D.C. Alley, N.F. 19920629:Tertiaryoccurrenceai the fern
i ygodiim(Schizaeaceae)inAustraliaandNewZealand.MenhirsoftheQueenslandMuseum
32(1): 203-222. Brisbane. ISSN0079-8835.
I.yyjhhum is ubiquitous in early Tertiary floras ofAustralia. Sterile and fertile leaves ofthe
climbing fern Lygodiumare found in Eocene deposits from Anglcsea(Victoria), Maslin Bay
and Golden Grove (South Australia) and in the early Tertiary from Dinmoie in Queensland
(L dinmorphylium Churchill. 1969). Sterile material is known from Late Eocene/Early
Oligocene sediments at Cethana (Tasmania) and the Middle Eocene at Temuka, in New
Zealand.
New material from theGolden Grove, Maslin Bay and Anglcsea localities including sterile
andfertilepinnules,spores,andcuticularstructureisreferredtoL.dinmorphylium.Thefertile
pinnae contain spores nfCyathidites splendens Harris, 1965. This palynomorph has a Laic
Cretaceous(Maastrichtian)toPliocene rangeinsoulhernAustralia, The-Australianspeciesis
very similar to the Eocene Chilean L skottsher^ii Halle, 1'MO, in sterile and fertile pinnule
morphology. Heterophylly presents problems in imerpretinrj variation in fossil Lygodium.
OLygodium, Schizaeaceae, Cyathidites splendens, Tetiu/rx, palytwtogy, pulaeabotuny,
biogeography,Australia, NewZealand.
AndrewC,Rozefelds, QueenslandMuseum,POBox3300,SouthBrisbane, Queensland4101,
Australia; present address: School of Botany, University of Melbourne, Parkvtlh 3052.
Victoria. Australia; D. C. Christophel BotanyDepartment, University ofAdelaide, POBox
496,Adelaide5001, SouthAustralia, Australia; N. F.Alley, SouthAustralian Departmentof
MinesandEnergy, POBox 151 Eastwood5053, SouthAustralia, Australia; 25June. 1991.
f
wiEtxhttaenmtpLeryagtoediouutmliiesrsaopcacnutrrroipnicgailncNleiwmbZienaglfaenrdn tHiolplheetlal(.1(917987)0).anDdimCghhr.iss(to19p7h8e)l, P&eteGrrse&enCwhroiosd-
I/..arricularumA.Rich.),easternUnitedStales(L (1987). Churchill's material was the first une-
pahiamm (Bernh ) Swartz and Japan [L quivocalfossilevidenceofthegenusinAustralia.
japonicum (Thumb.) Sw.]. About 40 species are The taxadescribedinthe 19th centurywere based
known world wide with four occurring in on fragmentary sterile pinnules but none can be
Australia(Holttum, 1959).TheAustralianspecies confidently assigned to Lygodium (Churchill.
L.japonicum(Thunb.)Sw.,L.flexuosum(L.)Sw.. 1969). Of these taxa the type of Osmunda tas-
/. tnurophylluni (Cav.) R. Br and L. rcticulatum mamca Johnston from Glenora, could not be lo-
Schkuhr arc tropical occurring in northern and cated incollections in Tasmania (R. S. Hill & N
eastern Australiu. In this paper, fossil Lygodium Kemp,pens.comm.).TheholotypcofL. strzeleck-
leaves including sterile and fertile pinnules are- ii Euingshausen, 1886, collected from the
described from Dinmore (Queensland), Maslin Vegetable Creek locality was restudied and
Bay and Golden Grove (South Australia) and prepared further (Fig. 3A) but referral to
Angiesea (Victoria) and a number of other Lygodium is dubious because ii is based upon
localities in southern Australia and New Zealand small pinnule fragments with abraded margins.
(Fig. I). Previous records of Lygodium or The Late Tnassic Lygodium antiquorum Shirley.
Lygodittm-tike:. ferns are reviewed. Problems in- 1898 from the Ipswich Coal Measures is based
herent in classifying and describing variable taxa uponatrilobateseedfernorfern pinnule(Fig 3F.i
arediscussed. withnoevidenceofsporangia,and isnotreferable
to Lygodiumas Walkom (1917) indicated.
PREVIOUS RECORDS OFFOSSIL Churchill (1969) described L dinmorphylium
LYGODWMINAUSTRALIA based upon two fertile pinnules from the early
AND NEW ZEALAND
Tertiary RedbankPlains Formationat Dinmorein
south eastern Queensland. Hill et al. (1970)
Lygodium or LygodiumAikc fossils have been recorded sterile Lygodium pinnules from Din-
reported by Johnston (1894), Ettineshausen more, which they referred to L. skottsbtrgii.
(1886. 1888), Shirley (1898). Churchill (1969), Douglas (1978) figureda fertile Lygodium pinna
204 MEMOIRS OFTHEQUEENSLANDMUSEUM
fromAngleseainVictoria. Afernfrondtentative- tion, south eastern Queensland is of
lycomparedwithLygodiumhasalsobeenfigured Paleocene/Eocene age (Day et al., 1983). This
from the Cenomanian Winton flora of Western largely undocumented flora includes ferns,
&
Queensland (Peters Christophel, 1978). Chris- Podocarpaceaeanddicotyledonousleaves(Hillet
tophel & Greenwood (1987) recorded Lygodium al., 1970; Selling, 1950).
from theGolden Grove locality. ThediverseAngleseafloraoccursintheMiddle
Additional material was collected from Din- Eocene Eastern View Coal Measures, and while
moreandAngleseabyACRandDCCrespective- Lygodiumoccursinmostlensesthemajoritycome
lyoveranumberofyears.Lygodiumhasalsobeen from the Gymnostoma lens (Christophel et al.,
recentlycollectedfromtheMiddleEoceneMaslin 1987). Elements of the Maslin Bay flora were
Bay and Golden Grove floras (South Australia) described by Blackburn (1981) from a car-
and the Late Eocene/Early Oligocene Cethana bonaceous clay lens in the Middle Eocene North
flora of Tasmania (R. Carpenter, pers. comm.). Maslin Sands (McGowran et al., 1970; Alley,
Lygodiumpinnaehavealsobeenrecognizedfrom 1987). A well preserved flora from the North
Temuka in New Zealand (D. Mildenhall, pers. Maslin SandsatGolden Grove, northofAdelaide
comm. 1986). is also Middle Eocene in age. A recent overview
&
of the flora has been provided by Christophel
LOCATION OFSITES AND AGES Greenwood (1987).
OFFLORAS
TheLateEocene/EarlyOligoceneCethanaflora
has yielded Nothofagus leaves (Hill, 1984), and
TheDinmoresiteintheRedbankPlainsForma- the remaining flora, which includes 'tropical'
LYGODIUM LOCALITIES
20
•1Dinm ore
• A/egetable Creek
30
40
Cethana
^<
5JGIenora
Temuka
FIG. 1. FossilLygodiumlocalities inAustraliaandNewZealand.
TERTIARYOCCURRENCEOFFERN LYGODTUM 205
FIG.2.LygodiumarticulatumA.Rich.fromFairyFallsTrack,WaitakereRange,SouthofAuckland,NewZealand.
DSIR,ChristchurchCHR153015.Noteabscissionnodesatbaseofsterilepinnulesandthevaryingdevelopment
oflaminabetweensorophores. [PhotographsuppliedbyBryonyMacMillan,DSIR,Christchurch,NewZealand].
206 MEMOIRS OFTHEQUEENSLANDMUSEUM
TERTIARYOCCURRENCEOFFERNLYGODIUM 207
rainforestelements is presently being studied (R. Lygodium material collected by ACR for this
Carpenter, pers. comm. 1987). The geology and study, will be donated to the Queensland Her-
f
straugraphieinformationon theTemukapotteries barium(BRI).
claypits in New Zealandindicates anEocene age
foi this flora (Wellman, 1953; Gain 1967). SYSTEMATICPALAEOBOTANY
Specimens from these two localities were not
availableforstudy. FamilySCH1ZAEACEAE
METHODS ANDPRESERVATION Lygodium Sw.
OF FOSSILS
The new Lygodium material from the va
The Dinmore.. Cethana and Vegetable Creek Australian localities is considered to be con-
materials are preserved as impressions in fine specific with L dmmorphylhtm Churchill from
grained mudstones lacking cuticle Latex peels of Dinmore in south eastern Queensland. An
the surface of sterile Lygodlum pinnules from emended diagnosis of this taxon including ittfoi
Dinmore, examined with a SEM, yielded no cel- raation on spores, cuticle and foliar heteromor-
eluvlearr,inSfoErMmateixoanm(iDn.aDtiilocnhero.fpefrrsa.gcmoemnmt.sjoHfotwh-e pcohviesrmedisthgeivgerno,ssasmoCrhuprhcohliolgl'ysediestcherifpetritoinleopnilny-
originalspecimendoshowsomecellularinforma- nules
tsipoenc(iFmiegn.sJa1rAetBp)r-esTehreveMdasaslicnomBparyesasnidonAsngolresiema- Lygodium dinmorphvllumChurchill, 1969
pressions while Lygodlum leaves and sorophores (Figs4-10)
rumGoldenGroveare mummified'.Themethod
I '
tor preparing 'mummified' leaves follows that of 196^LygodiumdinmorphvllumChurchill,pp.257-265,
Christophel (1984). TheGoldenGroveLygodium fig 2
leavesareusuallyopaqueandcouldnotbestudied 1970 Lygodium xkothbcrgii Halle, Hill etal, pi. C/Jl,
using the fluorescence technique of Friedrieh &. fig i.
Schaarschmidt (1977). Small areas ofvenation in
these specimens were illuminated using narrow Emended Diagnosis
heamsoflight fromcold light sources. Sterile pinnules asymmetrical, usually 2-1
The mummified sorophores and small frag- lobed, although unilobate, quadrilobate and pen
ments of leaf cuticle from Golden Grove *ctc talobate pinnules occur. Lateral lobes tend to he
clcaned withconcentrated HFacidfora few days longerthan centra! lobes. Pinnule base is cvncaic
and the sporangia were gently macerated to totruncate,apexroundedtoacute. Pinnulespctio-
release spores. Spores were studied and photo- late, lacking abscission node al pinnule ba&
graphed with aZeissPhotoruicroscope Hi.Frag- Venationpalmate,dichotnmouslybranchingfrom
ments of cuticle from the sorophores and leaves the central vein, on each lobe, two 10 three (I
wee washed in distilled water, airdried and sput- before reaching leafmargin. Leaflength variable,
ter coaled with platinum. Material was examined maximumlobelengihuboul 1 10mm-Pinnulemur*
with a JEOL ISM 840 scanning electron micro- gin variable, undulosc in larger specimens,
scopewithlanthanumhexaboridefilamentat5kv. smooth in small pinnae, rarely lobed. Pinnules
Lygodlumexamined inthispaper(Appendix A) probably hypostomatie. Anticlinal walls of
includes material from the Queensland Museum abaxial epidermis strongly sinuous, stomata
(QMF), Geological Survey of Queensland anomocyuc. Stomatal apparatus 30^m long, <29-
(GSQF), University of Queensland (UQF). 32pjn) and 27\unwide-(25-32|im).
MuseumofVictoria (NMVP),University ofTas- Fertile pinnules in branched axe* often Ittfc
mania (UTC), University of Adelaide (Maslin lamina, although lamina sometimes present as
Bay.UAS; GoldenGrove,UAG),andGeological narrow wing or flange at the base ofSOfopfa
Survey of New Zealand (GSNZ). Modern Individual sorophores variable in size, ranging
FIG. 3. A. Holoiype, LygodiumsirzeleckiiEuingshausen, from VegetableCk, GNSWF8820. X 3. B.C. Ceibann
ferns. B.fernaff.Lygodium. sterilepinnule UTC235. C BilobatesterileLygodiumpinnule UTC232, X 2. D.E
TcmUka Lygodium, New Zealand. D, GSNZ25/90. Xf 1.7. E, GSNZ25/92, X 2. F, Holotype. Lygodlwt
iin/u/uorum Shirley. GSQF143. x 3.5 If) Mildcnhall (DSlR.Ncw Zealand Geological Survey, Lower II n
and R. Carpenter (University of Tasmania) supplied photographs of the Temuka and Cethana specimen*
respectively].
-
208 MEMOIRSOFTHEQUEENSLANDMUSEUM
from about4.0-14.0mminlength,uften unitedin FOLIARHETEROMORPHISM IN EXTANT
LYGODJUM
groups of 2-4. Spores referable to Cyathidites
splendent Harris, 1965. Sinuosity of anticlinal
walls of epidermis on sorophore lamina no* as Foharheteromorphismistheproductionofmor-
pronounced asin sterile pinnaI phologically variable leaf types on the same in-
dividual(Eckenwalder, 1980).Changesinthesize
Remarks and shape of successive leaves can be due to
TheGoldenGroveandAnglescalocalitieshave hctcroblastic development (Wareing & Phillips,
yieldedLygodiumsorophorcscontainingCsplen 1982). This may occur in the transition from
dens spores. C. splendent is also .i common juvenile to adult leaves or may result from en-
palynomorph in the Maslin Bay paiynofloru. vironmental factors. hiLygodiutn,transition from
While spores have not been isolated from the sterile to fertile pinnae results in morphologically
Dinmore sorophorcs and the cuticle is poorly distinct pinnae. Pinnae intermediate in characteLr
kmantoewrnia,l itto Lseedminsmoprrpefheyrlalbulmetthoanraefpeerdt.1hedonubetw abrettiwceuelnatsutmerainledaLndpaflermtailteump.inTnuhleeslaomcicnuariinnthe
dfuiJsltyridbiusttiionnctoftaCx.ons.plBeansdeednsuipnonthetheearwliydeTesrptrieaaryd aafernredtiLlsetrppoiannlgnmlayaetdouifmmtoihsrepvehexirtcyanr(teFdisugp.cecei2d;easnMLda.ntachrheteipscitunelnarutlue&ms
tahnadttLhaeteDiCnrmeotraceesooursopihnoAruesstarlasloiahaitdsC.eesmpslelnidkeenlys Zavada, 1987, figs 9,11), The degree of lamina
spores. Lygodiutndinmnrphylhanisclosely com- reductionvariesinfertileleafletsoiL.articulatum
L (Fig. 2).
parable with sh/itshergiibut is considered dis- In Lygodiutn, there is great plasticity ofform in
tinctbaseduponsporesandcuticubrinformationL. sterile and fertile pinnae, and fusion of pinnules
Halle (1940: 259) noted that the spores of
occurs. Stfilile leafletsinexta^tLarricw/tftam arc
vkonsbergii are finely punctate oralmost smooth characteristically unilobate although fused bilob-
and hence differfrom C. splendens* atn pinnules also occur. Fusion of pinnules may
The only contra-indication that the Lygodiurn explain some variation in fossil material from
material from Australiamay include a numberof
Australia.
taxa is the variation in the angle of lower onter Aberrant pinnules with accessory lobing were
branching in thefertileaxesat differentlocalities. collected fromDinmore, Maslin Bay and Golden
In a relatively complete non-distorted specimen .• (Fig. 9G.FD and were also figured for L.
from Anglcsca (Fig. SD), the angle ofbranching katdjussi Heen 136! by Manchester & Zavada
is (I30-15C/*), which is similar to that ofmodern (1987). In modemL rericulatum fromnortheast-
L ariiadatum (Fig. 2). The Dinmore material ern Queensland similarly deformed leave* aUo
typically has acutely angled lower order branch- occur.
ing in the fertile pinnae, i.e. (75-85°] (Fig. 4A,
G.H). Relatively complete and undtstorted fertile FOSSILLYGODIUMMORPHOLOGY
axes have not been collected from Golden Grove
or Maslin Bay so the angle ofbranching in the Taxonomic treatments of extant Lygodium are
fertile axes from theselocalities is unknown.The basedupontheformofsterileandfertilepinnules,
systematic implications are difficult toassess be- spores, venationandmodeofbranching. Holttum
cause insufficient material has been collected (1959: 39) commented that 'there is so much
fromenough localitiestoassess variation. <nleafletformduetotheageoftheplant,
FIG.4.FertileLygodiumdinmorphyllum pinnules from Dinmore. A,QMF14465, X 1.9, sporangial head freeof
lamina. B. QMF14530. x 2, lamina around sporangial heads. C,D, GSQF12971 C, x2.«. D, Xl.0. Ej\
QMFI4361.E, X3.1.F, X 1.G,H, QMF12544. G, X 3.3. H, X 1.
FIG-5.SterileLygodiumdinmorphyllumpinnulesfromDinmore,illustratingvariationinpinnules, A,QMFI8035.
GB,SQQFM1F125937240.J.CQ,MQfMF14I346366.5A.llDn,atQuMrFal28si5/.He.. E, QMF15319. F, QMF14974. G, QMF14363. H. UQF32006. 1,
Fro ('. Sterile Lygodium dinmorphyllum pinnules from Dinmore, illustrating variation. A.QMF 18035. X 1 H,
OMF14809, X 2. C, UQF32006, X 1.4. D, QMH4363, X 3.5.
FIG 7. Fertile and sterile Lygodium dinmorphyllum pinnules from Maslin Bay. A,B, Sterile trilobate pinnule,
UAS7I5.A. X 2. B, x l.C,FertilepinnuleUAS3524, x S. D.E.Sterileunilobatepinnulewiihpoorlvdeveloped
secondary lobe, UAS2274.D, X 13.fi; -< 1.0. F.G. Bdobatc slcnle pmnule. UASI2U. F, X 1.9. G. X 1,0.
TERTIARYOCCURRENCEOFFERNLYGODIUM 209
Figure4
210 MEMOIRSOFTHEQUEENSLANDMUSEUM
Figure 5
TERTIARYOCCURRENCEOFFERN LYGODIUM 211
Figure6
212 MEMOIRS OFTHEQUEENSLANDMUSEUM
Figure7
UG