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Tertiary Climates

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Tertiary Climates 4 Palaeobotanical evidence for Tertiary climates D. R. GREENWOOD Climate is the primary framework within which 1985;Upchurch & Wolfe, 1987),the terms mega- plant populations grow, reproduce and, in the thermal, mesothermal and microthermal are used longer time frame, evolve. However, reconstruc- here to describe the temperature characteristics tions of palaeoclimateare often based on the of the vegetationof the main climatic zones.The marine record (e.g. Quilty, 1984 and Chapter 3, definitions of Nix (1982; Kershaw & Nix, 1988) this volume) or on computer modelling (Sloan & are used here," although Wolfe's (1979, 1985) Barron, 1990).The latter type, in particular,suf- vegetation classification uses slightly different fer from their dependenceon simplifiedscenarios definitions. and must be tested against the terrestrial This discussion is restricted to the Tertiary, palaeontologicalrecord. Individual species and since pre-Tertiary floras are dominated by plant whole plant communities are morphologicallyand groups with no or few modern analoguesand so physiologicallyadapted to their physical environ- palaeobotanical indices of climate are (as yet) ment, most stronglyto climate,and so plant mac- unreliable. Numerous accountsof the Tertiary cli- rofossils are a proxy record of past climates mate history of Australia basedon palaeobotanical (Wolfe, 1971, 1985; Upchurch & Wolfe, 1987). evidence have been published (Kemp, 1978, Severalstudies have indicated that plant macro- 1981;Christophel, 1981, 1988; Nix, 1982;Trus- fossils provide a potentially accurate record of well & Harris, 1982; Christophel & Greenwood, Tertiary terrestrial palaeotemperatures @olfe, 1989), in some casesas part of global accounts 1979,1990; Read et al., 1990). Interpretationof (Axelrod, 1984; Wolfe, 1985). Many of these the climatic signal preservedin plant fossil assem- interpretationshave been basedsolely on nearest- blages is dependent on understanding (1) how living-relative analogy(NLR) and provide qualita- plants (and vegetation)interact with climate, (2) tive reconstructions of climate. how plant fossil assemblageswere formed, and Many North American studies have used a (3) how these assemblagesrelate to the original strong correlation berween the proportion of vegetation. woody plant specieswith entire leaf margins and The use of terms such as tropical, subtropical mean annual temperature (MAT) in modern and temperate in palaeoclimaticdiscussions is mesic forests (leaf margin analysis,Wolfe, 1979, potentially conflusing,as these terms are rarely 1990), and other aspectsof foliar morphologY,to (lati- defined climatically and have geographical 'C * Modi{ied liom Nix (1982): mcgathermal, > 24 (MAT); "C; "C tudinal) connotations that are inapplicable for meso-mcgathcrmal intcrzone, 20-24 mcsothcrmal, l4-20 much of the Tertiary. Following Wolfe (1979, (14-24 "C if the interzoncis includcd); microthcrmal,< l4'C. l44l Palaeobotanicalevidence for Tertiarv climates 45 reconstructboth the regional distribution ofphysi- Francis, 1990) and epiphyllous ftttgt found on ognomically defined vegetationtypes and palaeo- leaf cuticle (Lange, 1978, 1981, 1982) are other climates for the Late Cretaceous and Tertiary sources of climatic information from plant fossil (e.g. Wolfe, 1985; Upchurch & Wolfe, 1987; assemblages.These approachesprovide quantitat- Parrish & Spicer, 1988). Wolfe (1971, 1979, ive estimates for critical components of climate 1985) has argued that foliar physiognomic that can be tested against estimates from the approachesare preferred to NLR methodologies, marine record and climatic models. primarily due to inaccurate systematictreatments for many Tertiary floras. However, foliar physiog- THE NATURE OF THE PLANT nomic analysisis used here accordingto the meth- MACROFOSSIL RECORD odolory proposed by Greenwood (l99la; Christophel & Greenwood, 1988, 1989),in com- Leaf accumulationsin streamsor lakes constitute bination with recent quantitative analysesof key a biassed,but neverthelessdetailed, record of the NLRs, of Tertiary taxa (Nix, 1982; Kershaw & parent vegetation. This bias results from a Nix, 1988;Read & Hope, 1990;Read et a1.,1990) number of processesacting on the plant parts (pri- to interpret early Tertiary macrofloras from Aus- marily leaves)from the time of abscissionor trau- tralia (Figure 4.1). Wood anatomy (Frakes & matic loss to their eventual entombment in sediment and lithification (Ferguson, 1985; 135o r5o" Spicer, 1989; Greenwood, l99la): 1. Deciduous and evergreentrees may have dif- ferent representationin an assemblageif leaf- fall is asynchronouswith periods of maximum sedimentation. 2. Different plant speciesmay haveleaves that are dehiscent or nondehiscent(e.9. many palms, herbs and forbs). Nondehiscent leavesare less olffio likely to be fossilisedthan are dehiscentleaves. 3. Plant organs may have varying capacities for transport from the plant body. For example, leaveswith a low weight per unit area tend to travel further in air than do denser leaves. 4. Leaves of individual species(e.9. sun versus shade leaves) and different species decay at different rates. 5. The varying productivity of the plants is sig- nificant, with canopy trees producing copious amounts of leaves,swamping the litter-fall. Fossil leaf accumulationstherefore reflect only a subset of the original surrounding vegetation, 135o 1soo and in most situations represent only the local Figure 4.1 Location map of AustralianTertiary macrofloras plant communities (Greenwood, 199| a). Further- discussedin the text, and modern thermal regimes(adapted more, smaller plant macrofossil accumulations from Nix, 1982;Christophel & Greenwood,1989): l, Stuart may represent geologicallyinstantaneous records Creek(silcrete floras); 2, Nelly Creek;3, GoldenGrove; 4, of past vegetation. Individual sedimentary facies Maslin Bay; 5, Nerriga;6, BacchusMarsh; 7, Anglesea;8, LatrobeValley (Yallourn & Monvell);9, Pioneer;10, Cethana; often reflect different components of the local ll, Monpeelyata. vegetational mosaic (see e.g. Christophel et al., 46 D. R. GREENWOOD Table 4.1. Foliar physiognoniccharacteistia of leaf-litterfrom modem Australion rainforest A. Mean "/" leaaes Leaf size "h Forest type" Micro- Noto- Mesophylls entire leaf margins CMVF 6-25 50-70 I l-39 90.0-98.3 CNVF 16-55 35-77 2-22 34.3-57.0 SI\IVF 59-85 14-36 0-ll 87.O-es.3(Q!d) 7.0-37.0(I,{s!v) MV-FF' 74-80 19-24 l-2 56.7-96.4 MFF' 90-96 4-10 0 1.0-13.0 NMF. 96-99 r-4 0 not available B. "/" taxa Leaf size Foresttype" Micro- Noto- Mesophylls LSI 7o entire leaf margins CMVF 1l-18 58-78 1l-30 47-s9 82.4-95.0 Cl\ryF 30-46 14-63 0-36 33-43 50.0-81.8 SNVF 40-75 25-50 0-20 13-38 84.2-e0.0(Qld) r43-2s.0 (NSIV) MFF r00 0 33.3-40.0 "CMVF, complexmesophyll vine forest;CNVF, complexnotophyll vine forest; SNVF, simplenotophyll vine forest;MV-FF, microphyllvine-fern forest;MFF' microphyllfern forest;LSI,leaf sizeindex (from Webb, 1959; Tracey, l98Z). ,Basedon a singlesite (four samples). "Basedon Macphailet al.,l99l, Table 3 (LakeDobson and Precipitous Bluff). Datafrom Greenwood,l99la (Table l, in part) and1992. 1987). However, stratigraphically correlated The present analysis uses a dataset derived macroflorasmay be separatedin time (as a record from modern Australian rainforest leaf-litter and of the original vegetation, and hence climate) by fluvially deposited beds (Christophel & Green- tens of thousands of vears. wood, 1987, 1988, 1989; Greenwood, 1992; Table 4.1). A multivariate foliar physiognomic analysisfor the Australian Middle Eocene macro- evidence of climate Palaeobotanical floras is in preparation (D.R. Greenwood,unpub- Terrestrial plant fossil assemblagesrecord local lished data), however, here temperature climate usually over small time intervals. Each characteristics are derived primarily from uni- plant assemblagerepresents a single datum of cli- variate comparisons(e.g. Figure 4.2). Other Aus- mate, with separate assemblagesfrom a single tralian Tertiary macrofloras are discussed flora providing information about local variation. qualitatively using leaf size and margin data only, However, continuous deposition of terrigenous as full data are not available.These analysesare sedimentswithin a single basin (e.9. the Latrobe supplemented by quantitative and qualitative esti- Valley)preserve climate records spanningmillions mates based on NLR analysesfrom Read et al- of years. (1990; Read, 1990) and Nix (19821,Kershaw & Palaeobotanical evidence for Tertiarv climates +7 25 U o qJ tr E20 0) 0) (g EI) C' 0,, 10 20 30 40 50 60 Leaf size index Figure 4.2 Relationship (least-squares regression) between leaf size index of forest-floor leaf-liner from Australian rainforests and mean annual temperature (MAT), r : 0.94. (Adapted from Greenwood, 1992.) Nix, 1988).Forest nomenclature and typologyin (LSI : (% microphylls+ 2/" notophylls+ 3% this discussionfollows those of Webb (1959, mesophylls 100)/2; Wolfe, 1978; Burnham, 1968), Tracey (1982; Table 4.1) and Wolfe 1989), and MAT (Greenwood,l99la, 1992; (re7e). Figure4.2).Mean leaf length(specimens) is also strongly correlated with MAT (Greenwood, r99Z\. Foliar physiognomy and climate The LSI of the Tertiary macrofloras (where Greenwood(1992; Greenwood& Christophel, applicable)has been used to calculatethe prevail- 1990, unpublished data) suggests that the ing MAT using a linear regressionequation from relationshipbenreen leaf margin type and MAT modern litter (Figure 4.2). These MAT values for Australianforests is different from, and more must be consideredto be minimum estimates;the
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