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Polar Wildfires and Conifer Serotiny During the Cretaceous Global Hothouse

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Polar Wildfires and Conifer Serotiny During the Cretaceous Global Hothouse Polar wildfires and conifer serotiny during the Cretaceous global hothouse Chris Mays1, David J. Cantrill2, and Joseph J. Bevitt3 1Department of Palaeobiology, Naturhistoriska riksmuseet, Box 50007, S-104 05 Stockholm, Sweden 2Royal Botanic Gardens Victoria, Private Bag 2000, South Yarra, VIC 3141, Australia 3Australian Nuclear Science and Technology Organisation, User Office, B3, New Illawarra Road, Lucas Heights, NSW 2234, Australia ABSTRACT Several highly effective fire-adaptive traits first evolved among modern plants during A CHATHAM ISLAND B (Rekohu) W PACIFIC 44°14’ the mid-Cretaceous, in response to the widespread wildfires promoted by anomalously high E N OCEAN PACIFIC 40° ZEALAND atmospheric oxygen (O2) and extreme temperatures. Serotiny, or long-term canopy seed CHATHAM OCEAN storage, is a fire-adaptive strategy common among plants living in fire-prone areas today, 300km ISLANDS but evidence of this strategy has been lacking from the fossil record. Deposits of abundant S CHATHAM RISE fossil charcoal from sedimentary successions of the Chatham Islands, New Zealand, record 44° B PITT ISLAND 176°E 180° 176°W Tupuangi wildfires in the south polar regions (75°–80°S) during the mid-Cretaceous (ca. 99–90 Ma). 0784 (Rangiaotea) Beach 0799 0815 Newly discovered fossil conifer reproductive structures were consistently associated with Waihere 0807 0811 44°16’S Bay 0812 these charcoal-rich deposits. The morphology and internal anatomy as revealed by neutron 0813 2 km tomography exhibit a range of serotiny-associated characters. Numerous related fossils from 0827 similar, contemporaneous deposits of the Northern Hemisphere suggest that serotiny was a 0814 176°14’W 176°12’ 176°10’ key adaptive strategy during the high-fire world of the Cretaceous. Figure 1. Fossil locality maps. A: Eastern Zea- landia; gray areas—emergent; gray line—2000 INTRODUCTION METHODS m isobath. B: Northern Pitt Island, Chatham Islands, with fossil localities recorded in this The mid-Cretaceous (Albian–Turonian; 112– study. All locality numbers have the locality 90 Ma) was one of the warmest intervals in geo- Sample Details prefix CH/f, as outlined in the text. Modified logical history. Paleoclimate records indicate Specimens were collected from Cenomanian from Mays et al. (2015, with permission). some of the highest oceanic temperatures known (99–94 Ma) Tupuangi Formation outcrops of in the marine realm (Friedrich et al., 2012), and Pitt Island. Photographs were taken using a warming was primarily at high latitudes, lead- Canon EOS 700D digital SLR camera. Mac- Organisation (ANSTO), Lucas Heights, New ing to a relatively low temperature difference eral data were collected by soaking >40 g of South Wales, Australia. Except for the primary between the tropics and the poles (Huber et al., sedimentary rock from each target horizon in scan parameters, which are included in Table 1995). These conditions were promoted by high 1% HCl (diluted with deionized water) for 7–14 DR3, all experimental setup details follow those CO2 levels, estimated from fossil plant proxies days, and gently sieving with water. Macerals in Mays et al. (2017). as ~2–5 times preindustrial levels (Barclay et 0.5–10 mm in diameter were sorted, dried at 60 al., 2010; Mays et al., 2015). High CO2 levels °C for 24 h, then weighed. Charcoalified (fusin- RESULTS and polar temperatures promoted productiv- ized) and noncharcoalified (coalified) wood cat- ity in mid-Cretaceous polar forests (Beerling egories follow those in Scott (2010). Collection Charcoal, Resin, and Plant Fossil Records and Osborne, 2002). Evidence from sedimen- and curation details, a discussion on taxonomic Sediment samples from fossiliferous hori- tary inertinite (fossil charcoal) indicates that the nomenclature, and comments on statistical anal- zons of the mid-Cretaceous Tupuangi Forma- 1 atmospheric oxygen concentration (pO2) was as yses are in the GSA Data Repository . Locality, tion revealed high abundances of fossil charcoal high as 29% during the mid-Cretaceous (mod- lithofacies, and maceral data are in Table DR1; (Fig. 3), indicative of intermittent wildfires. All ern ~21%), the highest levels of the past 250 fossil specimen details are in Table DR2. sediment samples yielded fossil resin (amber), m.y. (Bergman et al., 2004; Glasspool and Scott, with one exception (B1476); fossil resin pieces 2010), making wildfires far more likely to occur Neutron Tomography: Experimental Setup are as large as 1 cm in diameter. Bulk sediment (Brown et al., 2012). Computed tomography methods involve samples showed a statistically significant posi- The Tupuangi Formation was deposited in the collection of images, or radiographs, by tive correlation between the following two vari- a vast riverine-deltaic system within the south capturing radiation that has penetrated a tar- ables: (1) resin mass/total bulk sample mass and polar circle (75°–80°S; Mays, 2015) and crops get sample; in the case of neutron tomography, (2) charcoal mass/total bulk sample mass (Pear- out on Pitt Island in the Chatham Islands, east- neutrons are transmitted for this purpose. The son’s r = 0.463; p = 0.035; N = 21); this suggests ern Zealandia (Fig. 1). A fossil conifer seed cone neutron source for this study was the Open-Pool a temporal association between wildfires and scale (ovuliferous complex, OC, herein; sensu Australian Lightwater (OPAL) reactor housed at resin production and/or excretion. Escapa et al., 2016; Fig. 2) and associated sedi- the Australian Nuclear Science and Technology One conifer fossil, Tupuangi Protodammara mentary record from these strata indicate the herein (Fig. 2; additional details on specimen presence of fire-adapted floras at south-polar nomenclature in the GSA Data Repository), 1 GSA Data Repository item 2017382, supplemen- latitudes during the mid-Cretaceous, an interval tary images, tables and videos, are available online at was particularly common on horizons with of high O2, CO2, and some of the highest average http://www.geosociety.org/datarepository/2017/ or on high charcoal and resin content. These fossils global temperatures in Earth history. request from [email protected]. have features common to both Araucariaceae GEOLOGY, December 2017; v. 45; no. 12; p. 1119–1122 | Data Repository item 2017382 | https:// doi .org /10 .1130 /G39453 .1 | Published online 11 October 2017 ©GEOLOGY 2017 Geological | Volume Society 45 | ofNumber America. 12 For | www.gsapubs.org permission to copy, contact [email protected]. 1119 Figure 2. Tupuangi Protodam- mara. A: Abaxial fossil impression AC E F G with apical cusp and in situ resin delineating abaxial resin canals; arrow—abaxial ridge, PL1255B. B: Adaxial fossil impression. Black arrow shows adaxial ridge, white arrow shows arcuate crest, G PL1228. C: Neutron tomographic RNA reconstruction of compression High fossil, volume rendering, abax- 2 mm ial view. RNA—relative neutron 2 mm attenuation, in situ resin rep- resented by highest neutron Low attenuation; grid width on RNA 674 2 3 5 spectrum indicates relative 891 D 10 11 transparency, PL1231A. D: Mor- B 12 13 H phology and resin canal paths; 14 15 colored canals can be traced ? from pedicellate region to distal margin. Canals: red—apical, blue—medial, orange—lateral, ? PL1231A. E: Seed cones of the 2 mm serotinous species, Pinus bank- Resin canals Surface morphology siana, for comparison (Britton Abaxial Adaxial arcuate crest / Resin reservoirs and Brown, 1913). F–H: Sche- Adaxial Abaxial / adaxial ridge matic reconstruction of two Resin Inferred 2 mm Seed contact specimens of Tupuangi Proto- ? ? areas dammara on a partial mature seed cone. F: Distal view (form based on PL1244); vertical line indicates cross section in G. G: Syn-fire cross section; fire destroys the resin adhesion between the subtending ovuliferous complexes (OCs), promoted by the large subsurface resin reservoirs on the distal surface. Note that the adaxial reservoirs are laterally offset from the section. H: Post-fire cross section; the cone opens hygroscopically for seed release (analogous to Pinus; Dawson et al., 1997), which would likely facilitate OC detachment, but the timing and mechanism are uncertain (i.e., question mark). and Cupressaceae, but the following support a in part by several dissected compression fossils. the use of resin as a binding agent to hold the stronger affinity with the latter family: (1) three Several of these anatomically preserved fossil cones closed during dormancy (Ahlgren, 1974; seed scars (cf. one seed/ovule per OC for Arau- specimens had no surface expression and were Lamont et al., 1991). cariaceae; Farjon, 2010); (2) no discrete ovulif- only revealed and analyzed by neutron penetra- The morphological features of the fossils are erous scale (cf. Araucaria, Wollemia); and (3) an tion of their enclosing silicate matrix. consistent with the requirements for serotinous elongated pedicellate base. The combination of cones (He et al., 2016). The distal thickenings preserved characters indicates that these fos- DISCUSSION on both abaxial and adaxial surfaces would have sils represent a Cupressaceae stem group within provided an interlocking cone structure, serving the taxodioid Cupressaceae (sensu Rothwell et Fossil Evidence for Fire-Adaptive Traits to seal the cone margins until the seed release al., 2011), with close alliance to Taiwania, Ath- Fire-adaptive traits among plants are those stage. While these thickenings are likely homol- rotaxis, and several extinct taxa (see the Data that promote survival and/or reproductive
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