Mays & Cantrill

Alcheringa: An Australasian Journal of Palaeontology

ISSN: 0311-5518 (Print) 1752-0754 (Online) Journal homepage: http://www.tandfonline.com/loi/talc20

Protodammara reimatamoriori, a new species of conifer (Cupressaceae) from the Upper Cretaceous Tupuangi Formation, Chatham Islands, Zealandia

Chris Mays & David J. Cantrill

To cite this article: Chris Mays & David J. Cantrill (2018): Protodammara reimatamoriori, a new species of conifer (Cupressaceae) from the Upper Cretaceous Tupuangi Formation, Chatham Islands, Zealandia, Alcheringa: An Australasian Journal of Palaeontology, DOI: 10.1080/03115518.2017.1417478 To link to this article: https://doi.org/10.1080/03115518.2017.1417478

Published online: 04 Jan 2018.

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Download by: [Naturhistoriska Riksmuseum] Date: 05 January 2018, At: 02:26 Protodammara reimatamoriori, a new species of conifer (Cupressaceae) from the Upper Cretaceous Tupuangi Formation, Chatham Islands, Zealandia

CHRIS MAYS and DAVID J. CANTRILL

Mays, C. & Cantrill, D.J., January 2018. Protodammara reimatamoriori, a new species of conifer (Cupressaceae) from the Upper Cretaceous Tupuangi Formation, Chatham Islands, Zealandia. Alcheringa XXX,X–X. ISSN 0311-5518.

Isolated conifer female reproductive structures are common fossil elements from Cenomanian (ca 99–94 Ma) charcoal- and resin-rich beds of the Tupuangi Formation, Chatham Islands, southwest Pacific Ocean. Recent findings have proposed that these are the oldest fossil evidence of serotiny, a highly successful fire-adaptive reproductive strategy common among tree species living in fire-prone areas today. Herein, we systematically describe the external morphological and anatomical features of these fossils, by employing a combination of manual extraction and neutron tomography techniques. We propose a new species of conifer, Protodammara reimatamoriori, and a re-examination of fossil material of the Protodammara type species facilitated an emendation of the genus. Protodammara shares numerous features with extant Cunninghamia, Taiwania, Athrotaxis, and several extinct taxa of Cupressaceae, and is interpreted as an extinct lineage of the early-divergent ‘taxodioid Cupressaceae’ stem group.

Chris Mays [[email protected]] Department of Palaeobiology, Swedish Museum of Natural History, Frescativägen 40, Stockholm 114 18, Sweden; School of Earth, Atmosphere and Environment, Monash University, 9 Rainforest Walk, Clayton, VIC 3800, Australia; David J. Cantrill [[email protected]] Royal Botanic Gardens Victoria, Private Bag 2000, South Yarra, VIC 3141, Australia; School of BioSciences, University of Melbourne, Parkville, VIC 3010, Australia. Received 17.9.2017; revised 7.12.2017; accepted 12.12.2017.

Key words: Palaeobotany, ovuliferous complex, Cenomanian, ﬁre ecology, polar forest, taxonomy.

THE TUPUANGI FORMATION ﬂora of the Chatham Brown et al. 2012). This study seeks to establish a new Islands is characterized by a low diversity of ferns, species designation encompassing these organically pre- lycophytes and non-vascular plants, a high abundance served conifer fossil specimens. Comparable fossils and diversity of Gondwanan conifers (Araucariaceae, have been described for over 130 years from the Cupressaceae and Podocarpaceae), and subsidiary Northern Hemisphere (Heer 1882), but this is the ﬁrst Ginkgoales and angiosperms (Pole & Philippe 2010, such record from the Southern Hemisphere. Mays et al. 2015a, 2015b, 2017b). One form of fossil conifer ‘seed cone scale’ (ovuliferous complex or ‘OC’ herein; see ‘materials and methods’ section) was found Geological setting to be particularly common in layers with high charcoal

Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 The Tupuangi Formation is expressed in outcrops of the and fossil resin (amber) content. Both conventional Chatham Islands, southwest Pacific, an archipelago near manual preparation and non-destructive neutron tomog- the eastern margin of the largely submerged continent, raphy were employed to study the fossils, the latter Zealandia (Fig. 1). This continent, which also includes technique revealing the resin canals’ anatomy based on New Zealand, Lord Howe Island, New Caledonia and the anomalously high neutron attenuation of the in situ Campbell Island, is almost entirely under water today resin compared with the surrounding organic fossil (Mortimer et al. 2017), but was largely emergent remains and matrix. This constellation of evidence during the deposition of the Tupuangi Formation revealed an adaptive function of the resin in response to (Campbell et al. 1993). The Tupuangi Formation has regular fire during the mid-Cretaceous (Mays et al. been biostratigraphically correlated to the Ngaterian– 2017a). This interval likely had the highest proportions Mangaotanean New Zealand Chronostratigraphic stages of atmospheric oxygen since the late Palaeozoic (Mildenhall 1994, Mays & Stilwell 2013); these corre- (Bergman et al. 2004, Glasspool & Scott 2010), which spond to the Cenomanian–Turonian Global Chronos- would have promoted the frequency and intensity of tratigraphic stages (ca 99–90 Ma, Raine et al. 2015). wildfires around the globe (Belcher et al. 2010, The unit was deposited in a vast riverine-deltaic system (Campbell et al. 1993) within the south polar circle (75–80°S; Markwick et al. 2000). © 2018 Geological Society of Australia Inc., Australasian Palaeontologists https://doi.org/10.1080/03115518.2017.1417478 2 CHRIS MAYS AND DAVID J. CANTRILL ALCHERINGA Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018

Fig. 1. A, Map of eastern Zealandia, boxed area in Fig. 1B, grey areas = emergent, grey outline = 2000 m isobath. B, Map of the Chatham Islands region, boxed areas in Fig. 1C, D, grey areas = emergent. C, D, Generalized geological maps with fossil localities recorded in this study, all locality numbers have the locality preﬁx ‘CH/f’ as outlined in the text; C, Waihere Bay area, northwest Pitt Island; D, Tupuangi Beach area, northeast Pitt Island. Modiﬁed from Mays et al. (2015b) with permission.

Materials and methods outcrop succession at southern Tupuangi Beach Localities have GNS Fossil Record File numbers (preﬁx (Fig. 1). Stratigraphic heights of the localities corre- ‘CH/f’), and all (except CH/f0815) are from the spond to the outcrop successions illustrated by Mays & Waihere Bay outcrop succession; CH/f0815 is from an Stilwell (2013, Fig. 4 therein). Fossil material was ALCHERINGA PROTODAMMARA REIMATAMORIORI 3

collected by C.M., D.J.C., P. Viegas, and T. Ziegler dur- Order CUPRESSALES Link, 1831 ing January–February, 2016. All fossil materials are Family CUPRESSACEAE Gray, 1821 housed at GNS Science, Lower Hutt, New Zealand. Hand samples have been assigned unique sample regis- Protodammara Hollick & Jeffrey, 1906 emend. tration (‘GNS reg.’) numbers (prefixed ‘PL’), and Type species. Protodammara speciosa Hollick & Jeffrey, labelled with field numbers (prefixed ‘TPF’). In cases 1906. Lectotype (designated herein): 52818-1A where more than one fossil specimen was found associ- (Harvard University Herbaria); plate 1, Fig. 5 of Hollick ated with a hand sample, these specimens are provided & Jeffrey (1906). with letter codes (Table 1). Type locality. Kreischerville, Staten Island NY, Raritan Fifty-four fossil specimens (excluding seven speci- Formation, ca Turonian. men counterparts) are reported herein. These are preserved on or within 33 hand samples (PL1228–60); all Emended generic diagnosis. Ovuliferous cone scale specimens, except one, PL1235 (and its counterpart, complexes. Lignified, resinous. Spatulate in outline, PL1236A), can be ascribed to the newly erected species, pedicellate base, flattened, thickening distally. Bracts Protodammara reimatamoriori. All specimens are from have transverse ridge(s) subparallel to distal margin on strata of the Ngaterian and Arowhanan New Zealand one or both of the abaxial and adaxial surfaces. Ovulif- chronostratigraphic stages (Mays & Stilwell 2013), erous scale entirely fused with bract or otherwise indis- corresponding to the Cenomanian global chronostrati- tinct. Three seeds/ovules aligned in a single straight or graphic stage (ca 99–94 Ma, Raine et al. 2015). curved transverse row on the adaxial surface. Three or All except two compression and impression fossils more resin canals occur at the base of the ovuliferous were at least partly exposed by manual extraction; spec- complex; additional canals originate de novo or by imens PL1231G, PL1231H have no surface expressions, bifurcation. Variously developed apical process ener- and remain entirely encapsulated within sedimentary vated by a medial abaxial resin canal. matrix. Eight specimens were examined with neutron tomography (NT) from two hand samples (PL1231, Remarks. It was considered prudent to emend the gen- PL1244), using the experimental setup outlined by eric diagnosis slightly to accommodate the newly Mays et al. (2017b); the specific NT scan parameters described species, P. reimatamoriori, rather than erect a were presented by Mays et al. (2017a, table DR3 new genus for specimens of such close morphological therein). Resin volume estimates are based on the digi- and anatomical similarity to the type species. For a distal reconstructions of these eight specimens; the details cussion of these similarities, see the Discussion section. of estimating resin volumes from the neutron tomo- The genus was emended herein by: (1) removing the fi graphic reconstructions are outlined by Mays et al. size requirement, as this super cial character is more fi (2017a, supplemental materials therein). All resin canal typical of interspeci c, rather than intergeneric, variabil- dimensions and distributions were based on specimen ity; (2) specifying the number of resin canals that are PL1231A (Fig. 2C, F–O). present from the base, which was considered a better- fi ‘… fi Subclasses, orders and families follow the classifica- de ned character than the imprecise resin ducts ve ’ tion of Christenhusz et al. (2011), subfamilies follow or more, extending down the lower surface of the limb Gadek et al. (2000). All extant plant names adhere to the (Hollick & Jeffrey 1906, p. 199) with no indication of International Plant Names Index (IPNI Collaboration how far these canals extend; (3) including the termina- 2017). Synonymy symbols follow Matthews (1973). tion of a medial abaxial resin canal in the apical cusp; and (4) modernizing the descriptive terminology. Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 Character dimension ranges are presented in the following form: minimum (mean) maximum, N (where Protodammara reimatamoriori sp. nov. (Fig. 2) N = sample size); a full data-set of measured character dimensions is presented by Mays et al. (2017a, table v ? 2017a ‘Tupuangi Protodammara’; Mays et al., DR2 therein). The term ovuliferous complex (OC) pp. 1119–1122, Figs 2, DR2, videos DR1, DR2. employed herein refers to isolated seed-/ovule-bearing Diagnosis. Lignified ovuliferous complexes Spatulate organs without a clear distinction between bracts and outline with narrow, pedicellate base. Flattened, thicken- ovuliferous scales (Escapa et al. 2016); this term circum- ing distally but with thin, broad wings. Low-relief ridges vents any implications of homology or phylogeny. Resin on one or both abaxial and adaxial surfaces near, and sub- canals are provided with numbers starting with ‘1’ at the parallel to, the distal margin. A low-relief arcuate crest on midline; resin canal number increases with distance from the adaxial surface slightly proximal of adaxial ridge. the midline (even integers = left in abaxial and dorsal Abaxial surface weakly convex, unkeeled. Margin entire, views; odd integers = right in abaxial and dorsal views). except for a single apical cusp formed from a medial protrusion of bract. Ovuliferous scale completely fused with Systematic palaeontology bract. Seeds/ovules aligned on adaxial surface in a single transverse row. Three or more resin canals occur Subclass PINIDAE Cronquist et al., 1966 at the base; additional canals originate at or near the Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 HI ASADDVDJ CANTRILL J. DAVID AND MAYS CHRIS 4

Stratigraphic GNS Fossil Locality GNS Hand sample Sample Specimen Preservation Specimen Specimen height (m) Record File no. (FRF) ﬁeld no. Latitude (S) Longitude (W) reg. no. ﬁeld no. comments code style* type† notes

Waihere Bay 142 CH/f0811 WP39–5 m 44°15ʹ53.2ʺ 176°14ʹ17.5ʺ PL1228 TPF1244-01 A C R, O Paratype 1; Fig. 2B PL1229 TPF1521-01 Eight isolated A C R, O Fig. DR2A‡ specimens exposed B C R, O Fig. DR2A‡ C C R, O Fig. DR2A‡ D C R, O E C R, O F C R, O Fig. DR2A‡ G C R, O H C R, O Fig. DR2A‡ 147 CH/f0799 WP39 44°15ʹ53.2ʺ 176°14ʹ17.1ʺ PL1230 TPF1141B-06 Three isolated A C R, O specimens exposed B C R, O CI S PL1231 TPF1226-01 Six isolated A C R, O Paratype 2; specimens partly Fig. 2C, F–O; exposed, two Video DR1‡ entirely encapsulated within matrix B C R, O Fig. DR2C‡ C C R, O D C R, O EI S F C R, O G E R, O H E R, O 150.5 CH/f0807 WP38 44°15ʹ53.4ʺ 176°14ʹ16.6ʺ PL1232 TPF1632-01 A I S 155 CH/f0812 WP37 44°15ʹ53.5ʺ 176°14ʹ14.8ʺ PL1233 TPF1549-01 A I S PL1234 TPF1549-02 Counterpart to A C R, O PL1233 PL1235 TPF1553-01 A I S cf. Protodammara reimatamoriori PL1236 TPF1553-02 Counterpart to A C R, O cf. Protodammara PL1235; two reimatamoriori isolated specimens exposed B C R, O PL1237 TPF1631-01 A C R, O PL1238 TPF1706-04 A I R PL1239 TPF1708-01 A C R, O PL1240 TPF1708-02 A C R, O PL1241 TPF1708-03 A C R, O 286 CH/f0813 WP43 44°16ʹ23.8ʺ 176°14ʹ28.6ʺ PL1242 TPF1308A-01 Single, desiccated, A C R, O

isolated specimen ALCHERINGA PL1243 TPF1569-02 Two charred, A C R, O Fig. DR2B‡ isolated specimens exposed B C R, O PL1244 TPF1569-03 A C R, O Paratype 3; Fig. 2D, E; Video DR2‡ Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 LHRNAPOOAMR REIMATAMORIORI PROTODAMMARA ALCHERINGA 348 CH/f0827 WP46D 44°16ʹ24.1ʺ 176°14ʹ27.5ʺ PL1245 TPF1089A-01 A I R PL1246 TPF1089A-02 Counterpart to AI R PL1245 PL1247 TPF1089A-03 A C R, O PL1248 TPF1089A-04 Poorly preserved A C R, O PL1249 TPF1089A-05 Poorly preserved A I R PL1250 TPF1089A-05 Poorly preserved B C R, O PL1251 TPF1089A-06 Broken sample in AI R vial; very poorly preserved PL1252 TPF1089A-07 Poorly preserved A I R 360.5 CH/f0814 WP52 44°16ʹ26.7ʺ 176°14ʹ25.4ʺ PL1253 TPF1167-01 Three isolated A C R, O specimens exposed B C R, O C C R, O

Tupuangi Beach 251.5 CH/f0815 WP61 44°15ʹ47.9ʺ 176°10ʹ0.5ʺ PL1254 TPF1591-01 Three isolated A C R, O specimens exposed BI S C C R, O PL1255 TPF1591-02 Counterpart to AI S PL1254; two isolated specimens exposed B I S Holotype; Fig. 2A PL1256 TPF1594-01 A C R, O PL1257 TPF1594-02 Counterpart to AI R PL1256 PL1258 TPF1615-01 Two isolated A C R, O specimens exposed B C R, O PL1259 TPF1615-02 Counterpart to A C R, O PL1258; ﬁve isolated specimens exposed B C R, O C C R, O D C R, O E C R, O PL1260 TPF1635-01 A C R, O Table 1. Protodammara reimatamoriori sp. nov. specimen details. * (C)ompression; (E)ncapsulated in matrix; (I)mpression. † (R)esin; (O)vuliferous complex. ‡ Illustrations or animations presented by Mays et al. (2017a), but not included herein. 5 6 CHRIS MAYS AND DAVID J. CANTRILL ALCHERINGA

Fig. 2. Protodammara reimatamoriori sp. nov. fossil specimens, ab = abaxial, ad = adaxial, Fig. 2A, B, E–O reproduced from Mays et al. (2017a)

Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 with permission. A, Holotype, abaxial impression with apical cusp and in situ resin delineating some of the abaxial resin canals, arrow = abaxial ridge (PL1255B). B, Paratype 1, adaxial fossil impression, black arrow = adaxial ridge, white arrow = adaxial arcuate crest (PL1228). C, Paratype 2, partially exposed, desiccated compression fossil exhibiting in situ resin within the pedicellate base (PL1231A). D, E, Paratype 3, false colour surface renderings of a neutron tomographic reconstructed compression fossil (PL1244); D, Dorsal view, black arrow indicates apical cusp, note the concave adaxial surface; E, Oblique view of adaxial surface, arrows indicate interseminal ridges. F–O, Paratype 2 (PL1231A); F, neutron tomographic reconstruction, volume rendering, abaxial view, RNA = Relative Neutron Attenuation, in situ resin represented by highest neutron attenuation, grid width on RNA spectrum indicates relative transparency; G, Resin canal distributions, coloured canals are those that can be traced from pedicellate region to distal margin, canals: red = apical, blue = medial, orange = lateral; H, Outline sketch of Fig. 2F, G indicating section orientations of Fig. 2I–O; I–O, Volume renderings (J, L, N) employ the Relative Neutron Attenuation spectrum of Fig. 2F, line drawings (I, K, M, O) employ the canal numbers and colours of Fig. 2G, sections based on PL1231A, additional details from PL1244 (surface morphology) and PL1255B (canal distributions); I, Medial longitudinal section; J, K, Distal pedicellate transverse section, legend, scale and orientation apply to all transverse sections (J–O); L, M, Distomedial transverse section; N, O, Distal transverse section.

distal end of the pedicellate region de novo or via bifurca- Etymology. Moriori: reimata-, ‘tears of’ in reference tion to form two series: abaxial and adaxial. Distally, to the tear-shaped outline of an individual OC; -moriori, most canals arranged in a row close to abaxial surface; a the name and language of the original inhabitants of the smaller number arranged in a row close to adaxial sur- Chatham Islands where the fossils were found. face. The medial abaxial canal enters the apical cusp; two canals delineate the lateral wing margins; canals termi- Type material. Holotype: PL1255B (Fig. 2A), nate at or near abaxial ridge or distal margin. southern Tupuangi Beach, Chatham Islands. Tupuangi ALCHERINGA PROTODAMMARA REIMATAMORIORI 7

Formation, Cenomanian. Paratype 1: PL1228 (Fig. 2B); dominant, abaxial resin canals aligned in a single plane Paratype 2: PL1231A (Fig. 2C, F–O and video DR1 of (one ‘apical’ [no. 1]; two ‘medial’ [no. 6, no. 7]; Mays et al. 2017a); Paratype 3: PL1244 (Fig. 2D, E Fig. 2C, F–O). There are two smaller, ‘lateral’ canals and video DR2 of Mays et al. 2017a); paratypes are on the adaxial side of the main series (no. 14, 15); these from Waihere Bay, Chatham Islands. Tupuangi Forma- two canals of the adaxial series are evident near the tion, Cenomanian. base (Fig. 2A), but it is unclear if these diverge from the main series, arise from the cone axis or arise de novo. In the distal ‘winged region’ (Fig. 2F, G), up to Description 10 additional canals are present. Ovuliferous complexes preserved as lignified compres- The abaxial series includes: A, the apical canal (no. sions or impression fossils. All specimens are isolated; no 1), which enervates the apical cusp; B, the two medial cone axes with in situ OC specimens have yet been found. canals, which bifurcate at least once, resulting in four or more canals (no. 4–7) terminating in the vicinity of the abaxial ridge; C, two additional canals (no. 8, no. External morphology 9), which probably arise de novo, taper to distinct resin- fi Outline spatulate and slightly asymmetrical (across longi- lled tips and terminate at the distal margin approxi- tudinal axis). Ovuliferous complexs with maximum pre- mately halfway between the apical cusp and each lateral – served thickness of 1.5 (1.8) 1.9 mm (N = 7) within the wing margin (Fig. 2F O); and D, approximately four – distalmost 25% of the longitudinal axis length. Total small canals of uncertain origins (no. 10 13), which ter- length: 6.5 (8.7) 11.4 mm (N = 24). Proximally, OC char- minate close to the lateral margins (between no. 8 and acterized by a narrow, elongate pedicellate base with sub- no. 14 or no. 9 and no. 15). The adaxial series includes: parallel sides, straight to weakly convex basal margin and A, the two lateral canals (no. 14, no. 15), which follow oval to rhombic cross-section; ‘pedicellate region’ diame- the arced lateral margins of each of the OC wings, ter- ter: 1.4 (2.2) 3.4 mm (N = 25). Uncommonly, pedicellate minating at the edges of the distal surface; and B, two region thinner in the medial section, forming concave lat- large adaxial canals (no. 2, no. 3), which might bifur- eral margins. Distally, OC broadens into a ‘winged cate from the apical canal or originate de novo, are region’ characterized by two thin lateral wings; diameter proximal to the apical canal, and terminate near the of ‘winged region’: 6.2 (8.5) 12.7 mm (N = 29). Distal adaxial ridge (Fig. 2G, I). OC margin convex and entire, except for a single apical Resin canals are round or elliptical in transverse sec- cusp 0.3 (0.53) 0.93 mm long (N = 12) formed from a tion, with the exceptions of no. 2 and no. 3, which are distomedial protrusion of the bract. Bract intercalary crescentic near the distal margin (Fig. 2N, O). All growth represented by: (1) narrow, low-relief transverse canals enlarge distally, generally reaching maximum ridges on the abaxial (Fig. 2A) and adaxial (Fig. 2B) sur- width within the distalmost 25% of the OC. The largest – faces close and subparallel to the distal margin, maximum canals (no. 1 3) are up to 0.9 mm in diameter. distance from distal margin: abaxial 0.79 (0.96) 1.13 mm Estimated resin volumes per OC are 17.1 (30.4) 41.9% (N = 4); adaxial 0.49 (0.61) 0.72 mm (N = 7); in some (N = 8), as a proportion of total OC volume (see table cases, one or both of these can be quite reduced (e.g., DR4 of Mays et al. 2017a). Fig. 2D, E); and (2) a single arcuate crest slightly proximal of the adaxial ridge (Fig. 2B, E). With the exceptions of the ridges and crest outlined above, the distal region of Remarks Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 the OC has a weakly concave to very weakly convex The ovuliferous scale and bract are entirely fused in adaxial surface and weakly convex abaxial surface Protodammara reimatamoriori, a common feature in (Fig. 2D). These indicate small (max. length: 1.95 mm; several taxodioid Cupressaceae groups. Taiwania max. width: 1.17 mm) ovoid to reinform, probably abor- Hayata, Athrotaxis D. Don and Sequoioideae exhibit the tive ovules. Two weakly expressed, longitudinal inter- complete fusion of the ovuliferous scale and bract as seminal ridges are preserved on the distomedial adaxial they develop, whereas Cunninghamia R. Br. in Rich. surface of some specimens; adjacent to these ridges are consistently retains a distinct remnant of the scale even three oval, longitudinally aligned seed scars (Fig. 2E). at maturity (Farjon & Ortiz Garcia 2003). In taxa where One specimen identified with three seed/ovule impres- this fusion occurs, the mature seeds are anatropous, and sions on the adaxial surface near the distal end of the aligned longitudinally on the medial portion of the pedicellate region (PL1228). There was no evidence of a adaxial surface (Schulz & Stützel 2007). Two longitudi- distinct ovuliferous scale on any observed specimens. nal interseminal ridges proximal of the arcuate crest indicate three seed scars (Fig. 2E). This supports the hypothesis that the arcuate crest delineates the distal Resin canal anatomy margin of a fused/adnate ovuliferous scale, but further All resin canals are part of either an abaxial or an adax- evidence is required to establish whether these seeds ial series. At the base of the OC, there are three had anatropous orientations. 8 CHRIS MAYS AND DAVID J. CANTRILL ALCHERINGA Discussion & Sato 2007), has similar gross morphology to P. reimatamoriori, but lacks an apical process. It is Comparisons: Protodammara and ‘Dammara’ probable that a systematic re-examination of one or ‘ ’ Similar isolated OCs have been described from Creta- more of the above variants of Dammara , perhaps with ceous strata of the Northern Hemisphere for over neutron or other tomographic methods, will result in 130 years. They have generally been assigned to ‘Dam- several of these being validly assigned to the newly mara’ Lamarck or Protodammara (Heer, 1882, White, emended Protodammara. 1890, Hollick, 1892, 1897, 1898, 1902, 1904, Newberry, 1895, Knowlton, 1905, Hollick & Jeffrey, Comparisons: Araucariaceae vs Cupressaceae 1906, 1909, Kimura & Sekido, 1978). The present specimens represent the first record of similar OCs in the As with the OCs referred to as ‘Dammara’ (a junior Southern Hemisphere. ‘Dammara’ was never validly synonym of Agathis; Araucariaceae) mentioned above, published, and is a junior synonym of Agathis Salisb. the name Protodammara reflects the araucariacean (Bakhuizen van den Brink 1955, Grimaldi et al. 2000). affinity initially interpreted for this genus (Hollick & Protodammara was erected to describe mid-Cretaceous Jeffrey 1906, 1909). However, critical reviews of these fossils from northeastern USA that were similar to, but taxa have since concluded that both Protodammara and smaller than, most of the previously known ‘Dammara’ ‘Dammara’ have features typical of taxodioid Cupres- scales. When compared with the type material, and the saceae. Miller (1977) drew unspecified similarities of additional specimens of Hollick & Jeffrey (1909), the Protodammara (and ‘Dammara’)toCunninghamia, the presently described specimens share important charac- only extant genus of Cunninghamioideae (Cupres- ters with the type species, Protodammara speciosa (ca saceae); no taxonomic justification for this comparison Turonian; Grimaldi et al. 1989). These characters was made at the time, but Miller supported this asser- include: at least three basal resin canals, an apical cusp, tion by invoking a similar conclusion purportedly made which is fed by an abaxial resin canal, distal thickening by Krassilov (1971). However, Krassilov did not pro- expressed as abaxial and adaxial transverse ridges, three vide a taxonomic comparison, and made no mention of seed/ovules and a pedicellate-spatulate outline. Cunninghamia in reference to the fossil taxon. Later, Protodammara reimatamoriori can be distinguished Miller (pers. comm. in Grimaldi et al. 2000) expanded from P. speciosa by having an adaxial arcuate ridge on his cupressaceous interpretations for ‘Dammara’ and and broad, thin distal wings; it is worth noting that Protodammara, making the inference that the large P. reimatamoriori is consistently larger, but size is not a bract fused with the ovuliferous scales were indicative diagnostic criterion. of Cupressaceae. This character is insufficient to distin- No species of ‘Dammara’ has been thoroughly guish these OCs from various members of Araucari- described anatomically described, so comparisons must aceae, e.g., Agathis (Farjon, 2010). Direct-Temperature rely on gross morphological features at present. Proto- Resolved Mass Spectrometry was performed on the dammara differs from ‘Dammara’ microlepis Heer, in situ resin of ‘Dammara’-like scales from the Raritan 1882, and ‘Dammara’ borealis Heer, 1882 (both from Formation (Turonian); the spectral signature revealed Cretaceous strata of Greenland), by lacking the similarities to Pinaceae, but Cupressaceae could not be distinctive longitudinal, arcuate grooves of these taxa. ruled out (Grimaldi et al. 2000). As it stands, there has Protodammara is consistently smaller, but this is a been inadequate evidence put forth to categorically non-diagnostic character of the newly emended genus. assign all taxa of ‘Dammara’ and Protodammara to Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 Furthermore, P. reimatamoriori has three distomedial one family. Rather, we recommend each taxon be seed scars, rather than the single basally positioned scar reviewed individually, as we have done for Proto- of ‘Dammara’ borealis. The present taxon differs from dammara below, before reliable affinities can be inter- ‘Dammara’ cliffwoodensis Hollick, 1897 (Magothy preted for any isolated ‘Dammara’-like OC specimens. Formation, Turonian–Santonian; Christopher 1979), by The following combination of characters support an having a shorter pedicellate region and a broader distal affinity with Cupressaceae rather than Araucariaceae (or winged region, and from ‘Dammara’ northportensis Pinaceae) for Protodammara: (1) three elongate seed Hollick, 1904 (ca Turonian; Grimaldi et al. 2000), by scars, whereas extant Araucariaceae have a single seed having a distinctly convex and cuspidate distal margin per OC (Farjon 2010); although anomalous cases of rather than being incurved/concave. ‘Dammara’ acicu- multiovulate OCs have been reported (e.g., Araucaria laris Knowlton, 1905, of the Judith River Formation, sect. Bunya), there have been no fossils of confirmed Montana (ca Campanian; Goodwin & Deino 1989), has araucariacean affinity consistently exhibiting multiovu- a much longer apical process than P. reimatamoriori, late OCs (Dettmann et al. 2012, and references therein); and features several longitudinal ridges, which the latter (2) a lack of discernible ovuliferous scale or associated taxon lacks. Protodammara sp., an isolated OC from ‘ligule’,asperAraucaria de Jussieu (Wilde & Eames Lower Cretaceous strata of central Honshu, Japan 1952)andWollemia Jones et al., 1995 (Jones et al. (Kimura & Sekido 1978; ca Barremian–Aptian; Umetsu 1995, Dettmann et al. 2012); and (3) a pedicellate base, ALCHERINGA PROTODAMMARA REIMATAMORIORI 9

in contrast to the cuneate base typical of Araucariaceae. Cunninghamiostrobus goedertii and C. yubariensis Furthermore, this species shares distinctive similarities (sensu Stopes & Fujii 1910) have comparable canal and probable homologies with several Cupressaceae widths and distributions to the present taxon, but members, as explored below. C. goedertii, C. hueberii and C. yubariensis all have evidence of distinct ovuliferous scales with free margins and/or apices. The complete fusion of the ovuliferous Comparisons: Extinct and extant taxodioid scale and bract in the Protodammara precludes designa- Cupressaceae tion to Cunninghamiostrobus. There are important morphological differences Protodammara exhibits numerous characters in com- between the type specimen of C. yubariensis described ‘ ’ mon with taxodioid Cupressaceae (sensu Rothwell by Stopes & Fujii (1910), and Cunninghamiostrobus fi et al. 2011), which is a paraphyletic group de ned as yubariensis sensu Ohana & Kimura (1995; — Cupressaceae (Callitroideae+Cupressoideae). Of par- ‘C. yubariensis’ herein). In addition, the emended diag- ticular interest in this regard are: (1) Cunninghamia, nosis of Cunninghamiostrobus by Ohana & Kimura Taiwania and Athrotaxis, which are generally consid- (1995) is rejected here because of its redundancy with ered the most basal extant genera of Cupressaceae in Cunninghamia. This is exemplified by the following phylogenetic analyses (Gadek et al. 2000, Schulz & statement: ‘cone scales (OC herein) and axis in structure Stützel 2007, Mao et al. 2012); and (2) several early- like those of the living genus Cunninghamia’ (p. 140); divergent and extinct taxa representing cupressaceous the remaining characters in the diagnosis do not allow stem groups. These similarities are expanded on below, for adequate distinction from Cunninghamia. Features with additional comparisons made to approximately of ‘C. yubariensis’ which do not conform to Cunning- coeval eastern Gondwanan (Australia and Zealandia) hamiostrobus sensu Stopes & Fujii (1910) include: (1) members of the Sequoioideae. A summary of compar- a complete fusion of ovuliferous scale and bract; (2) a isons between these fossils and various extinct and small number of simple resin canals; and (3) a greater extant taxa of Cupressaceae was presented by Mays degree of bract intercalary growth reflected by an abax- et al. (2017a, table DR5 therein). ial ridge and ‘adaxial swelling’. These differences sug- The OCs of Cunninghamia are foliate, coriaceous, gest that this specimen should not be placed in with acuminate apices and a reduced but distinct ovulif- Cunninghamiostrobus. The adaxial swelling is similar erous scale on the adaxial surface of each bract. The in general form to the arcuate crest of Protodammara fi ligni ed structure, prominent adaxial and abaxial ridges reimatamoriori, and this swelling exhibits ovular attach- and lack of distinct ovuliferous scale differentiate Proto- ment points (Ohana & Kimura 1995). These attachment dammara from Cunninghamia, despite several impor- points were not observed in P. reimatamoriori, but both tant shared characters, such as having numerous abaxial species share the interseminal ridges between three seed resin canals, a similar number of ovules/seeds and a scars. Another similarity between ‘C. yubariensis’ and pedicellate base. Well-preserved Cretaceous cunning- Protodammara includes the abaxial ridge (Ohana & hamioid ovulate cone taxa have been recorded from Kimura 1995); in addition to the adaxial swelling, this Canada: the Early Cretaceous monotypic Hubbardias- ridge probably represents a degree of bract intercalary trobus Atkinson et al., 2014a; and three Late Creta- growth common to both taxa. Finally, an indistinct ceous taxa, Acanthostrobus Klymiuk et al., 2015, ovuliferous scale occurs in both ‘C. yubariensis’ and Cunninghamia taylorii Serbet et al., 2013, and Hugh- Protodammara. These comparisons suggest a close rela-

Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 millerites vancouverensis Atkinson et al., 2014b.In tionship, but the differences in gross morphology, resin fi addition to these, a coali ed Cunninghamia-like seed canal distribution and a lack of bract keel in the present cone, Pentakonos diminutus Herrera et al., 2017, has species preclude conspecificity. However, these features been reported recently from Lower Cretaceous strata of of ‘C. yubariensis’ are compatible with the emended fi Mongolia. These ve taxa have membranous tips or diagnosis of Protodammara; a re-examination of the arcs on the adaxial surface, which delineate the margins ‘C. yubariensis’ material is warranted to support this of the ovuliferous scale. Similarly, Elatides Heer, 1876 redesignation. emend. Harris 1979, has a similar gross morphology to Taiwanioideae is represented by a single extant spe- Protodammara reimatamoriori, but has a membranous cies, Taiwania cryptomerioides Hayata, and a sparse ‘ ’ ligule on the adaxial surface, which is likely an record of Taiwania-like fossils since the Jurassic. A expression of the ovuliferous scale. In contrast, the recent phylogenetic analysis of morphological and ovuliferous scale of Protodammara is entirely fused anatomical characters from a range of extinct and extant with, or otherwise indistinguishable from, the early-divergent Cupressaceae (Herrera et al. 2017) placed bract. Cunninghamiostrobus Stopes & Fujii, 1910, has Stutzeliastrobus foliatus Herrera et al., 2017, as a sister been recovered from Cretaceous strata of Japan group to a polytomy between: (1) Taiwania; (2) (C. yubariensis Stopes & Fujii, 1910) and California Austrohamia minuta Escapa et al., 2008; and (3) (C. hueberii Miller, 1975), and Oligocene strata of a crown group consisting of Athrotaxidoideae+ Washington (C. goedertii Miller & Crabtree, 1989). Sequoioideae+Cupressoideae+Callitroideae. Protodammara 10 CHRIS MAYS AND DAVID J. CANTRILL ALCHERINGA

reimatamoriori shares numerous features with Taiwania sion preservation style of these specimens. Athrotaxites and Taiwania-like fossil taxa, such as S. foliatus and A. min- stockeyii Escapa et al., 2016, was described from iso- uta, so an in-depth comparison is warranted here. The OCs lated, fertile athrotaxidoid OCs, and their preserved of Taiwania cryptomerioides (and the possibly conspecific internal anatomy reveals both abaxial and adaxial resin Cretaceous fossils from Alaska reported by LePage 2009) canals of similar size and number to those of P. have the following features in common with P. reimatamor- reimatamoriori. Furthermore, although the bract thick- iori: no distinct ovuliferous scale, thin broad wings in the ening is substantially greater than P. reimatamoriori, A. distal region, a system of branching resin canals within the stockeyii has a distinct adaxial arcuate crest and abaxial bract, and a variably developed adaxial arcuate crest. How- ridge. However, A. stockeyii differs from P. reimatamor- ever, the OCs of T. cryptomerioides are distinctly foliate iori by possessing non-bifurcating resin canals and the with minimal intercalary growth, and produce a maximum prominent acuminate apex characteristic of other of two seeds per OC. The OCs of Stutzeliastrobus foliatus members of Athrotaxites. from the Lower Cretaceous of Mongolia have no distinct Extant members of Sequoioideae are known only ovuliferous scale and a similar outline to P. reimatamoriori, from the Northern Hemisphere; however, one fossil- but differ from the present species by having minimal bract genus of this subfamily, Austrosequoia Peters & Christo- thickening, and having only a single resin canal. Austro- phel, 1978, has been recorded only from mid-Cretaceous hamia minuta is the oldest known probable member of to Oligocene strata of eastern Gondwana (Ettingshausen Cupressaceae (Early Jurassic; Cúneo et al. 2013); the OCs 1887, Peters & Christophel 1978, Hill et al. 1993, of this taxon have no discrete ovuliferous scale, similar to McLoughlin et al. 1995, 2010, Mays et al. 2017b). Two P. reimatamoriori. Key features that differentiate this spe- species of Austrosequoia are approximately coeval with cies from P. reimatamoriori include a distinctive abaxial Protodammara reimatamoriori: A. novae-zeelandiae keel and, like Stutzeliastrobus foliatus, a foliate structure Ettingshausen, 1887 (Zealandia; Mays et al. 2017b), and with minimal (if any) bract thickening. Candidates for A. wintonensis Peters & Christophel, 1978 (northeastern extinct taiwanioid groups are the monotypic genera Paratai- Australia). Morphologically, the OCs of Austrosequoia wania Nishida et al., 1992,andMikasastrobus Saiki & differ from Protodammara by having prominent distal Kimura, 1993. Parataiwania nihongii Nishida et al., 1992, intercalary growth and truncated apices; these features has a very similar resin canal distribution to P. reimatamori- are characteristic of all known members Sequoioideae. ori, but has only a single basal canal, in contrast to at least Furthermore, anatomically preserved silicified specimens three for P. reimatamoriori. Furthermore, both Parataiwa- of the type species, Austrosequoia wintonensis, have a nia and Mikasastrobus had a thin but conspicuous ovulifer- greater number of seeds (four to seven) and small, trans- ous scale; there is no evidence of this in Protodammara. versely arranged resin bodies (vs prominent, radially Athrotaxidoideae is represented by a single extant arranged resin canals in Protodammara). genus, Athrotaxis, endemic to Tasmania, Australia. Like There are numerous taxodioid Cupressaceae taxa Protodammara reimatamoriori, the OCs of Athrotaxis that have eluded satisfactory classification, and share are woody and have no discernible ovuliferous scale; important characters with Protodammara reimatamori- however, Athrotaxis differs from the presently described ori. Sphenolepis Schenk, 1871, emend. Harris 1953, taxon by having a much greater degree of intercalary and the present species both have a pedicellate base, a growth resulting in a peltate to clavate form, and long, distally spatulate form, a cuspidate apex, an arcuate curved acuminate apical processes. There have been adaxial ridge and at least three basal resin canals. How- numerous reports of athrotaxidoid fossils since the Late ever, the peltate OCs of Sphenolepis are the result of a

Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 Jurassic (Dong et al. 2014), but reviews of Athrotaxis- greater degree of intercalary growth than P. reimatamor- like fossils have concluded that many such fossils have iori, reminiscent of Athrotaxis or Sequoia Endl., and dubious afﬁnities to Athrotaxidoideae; Florin (1960) possess two rows of seeds, whereas the present species concluded that there were no reliable occurrences of has a single row. Furthermore, the ovulate cones of Athrotaxis in the Northern Hemisphere, a view that has Sphenolepis (e.g., S. kurriana [Dunker, 1846] Schenk, since largely been supported (Miller & LaPasha 1983; 1871, emend. Harris, 1953; ca Aptian–Albian; Yans Dong et al. 2014) with the possible exception of et al., 2002) tend not to disintegrate, whereas the OCs A. parvistrobili Miller & Hickey, 2010, from Lower of P. reimatamoriori have never been found attached to Cretaceous (Albian) strata of Washington, USA. Despite their axes, suggesting typical shedding of OCs after this, there remain several probable athrotaxidoid taxa of maturation in the latter taxon. The probable cupressa- Cretaceous age reported from both hemispheres. Athro- ceous (or pinaceous) Pararaucaria Bock, 1954 (‘Para- taxites berryii Bell, 1956 emend. Miller & LaPasha raucariaceae’ sensu Taylor et al. 2009), has a dominant 1983, Athrotaxites yumenensis Dong & Sun, 2014, ovuliferous scale (Stockey 1977), which is absent in Athrotaxis parvistrobili and Athrotaxis ungerii Protodammara. Two ancient cupressaceous taxa have Archangelsky, 1963, all differ from Protodammara by recently been described from Scotland: Hughmillerites having large, curved acuminate apices. Thorough juddii (Seward & Bancroft, 1913) Rothwell et al. 2011 anatomical comparisons are precluded by the compres- (Late Jurassic), and Scitistrobus duncaanensis Spencer, ALCHERINGA PROTODAMMARA REIMATAMORIORI 11

Mapes, Hilton & Rothwell, 2015 (Middle Jurassic). Lower Cretaceous diversification of cunninghamioid Cupres- – These taxa differ from Protodammara by having foliate saceae. International Journal of Plant Science 175, 256 269. fi ATKINSON, B.A., ROTHWELL, G.W. & STOCKEY, R.A., 2014b. Hughmil- OCs lacking any signi cant bract thickening, and lerites vancouverensis sp. nov. and the Cretaceous diversification prominent ovuliferous scales. of Cupressaceae. American Journal of Botany 101, 2136–2147. In summary, despite a range of characters shared with BELCHER, C.M., YEARSLEY, J.M., HADDEN, R.M., MCELWAIN, J.C. & fl ’ several early-divergent Cupressaceae subfamilies, Proto- REIN, G., 2010. Baseline intrinsic ammability of Earth s ecosys- tems estimated from paleoatmospheric oxygen over the past 350 dammara lacks: (1) a distinct ovuliferous scale million years. Proceedings of the National Academy of Sciences (cf Cunninghamioideae); (2) a foliate form (cf. Cunning- of the United States of America 107, 22448–22453. hamioideae, Taiwanioideae); (3) a long apical process BELL, W.A., 1956. Lower Cretaceous floras of western Canada. Mem- – (Athrotaxidoideae); and (4) prominent intercalary growth oirs of the Geological Survey of Canada 285,1 331. BERGMAN, N.M., LENTON, T.M. & WATSON, A.J., 2004. COPSE: A new (cf. Athrotaxidoideae, Sequoioideae, Taxodioideae). This model of biogeochemical cycling over Phanerozoic time. Ameri- precludes designation of Protodammara to any extant sub- can Journal of Science 304, 397–437. family at present. Instead, this genus likely represents a BOCK,W.,1954. Primaraucaria, a new araucarian genus from the Vir- – distinct and extinct member of the ‘taxodioid Cupres- ginia Triassic. Journal of Paleontology 28,32 42. BAKHUIZEN VAN DEN BRINK, R.C., 1955. Nomenclatural note on Dam- saceae’ stem group. A re-examination of the numerous pre- mara Lmk and Agathis Salisb. Taxon 4, 195–196. viously reported Protodammara and Protodammara-like BROWN, S.A.E., SCOTT, A.C., GLASSPOOL, I.J. & COLLINSON, M.E., taxa of isolated OC fossils, particularly with NT, will likely 2012. Cretaceous wildfires and their impact on the Earth system. Cretaceous Research 36, 162–190. reveal: (1) their close phylogenetic placement with P. CAMPBELL, H.J., ANDREWS, P.B., BEU, A.G., MAXWELL, P.A., EDWARDS, reimatamoriori; (2) similar serotiny-related morphological A.R., LAIRD, M.G., HORNIBROOK, N.D.B., MILDENHALL, D.C., WAT- and anatomical characters; and (3) their relatively high TERS, W.A., BUCKERIDGE, J.S., LEE, D.E., STRONG, C.P., WILSON, diversity and widespread distribution concurrent with the G.J. & HAYWARD, B.W., 1993. Cretaceous-Cenozoic Geology and fi Biostratigraphy of the Chatham Islands, New Zealand. Institute re-prone climates of the mid-Cretaceous. of Geological and Nuclear Sciences Limited, Lower Hutt, New Zealand, 269 pp. CHRISTENHUSZ, M.J.M., REVEAL, J.L., FARJON, A., GARDNER, M.F., MILL, Acknowledgements R.R. & CHASE, M.W., 2011. A new classification and linear sequence of extant gymnosperms. Phytotaxa 19,55–70. Assistance with fossil preparation was provided by CHRISTOPHER, R.A., 1979. Normapolles and triporate pollen assem- members of the Monash Palaeontology Undergraduate blages from the Raritan and Magothy Formations (Upper Creta- Volunteer Program, with special thanks to Wayne Mays ceous) of New Jersey. Palynology 3,73–121. and Chief Preparator Chava Rodriguez. Field assistance CRONQUIST, A., TAKHTAJAN,A.&ZIMMERMANN,W.,1966. On the higher taxa of Embryobionta. Taxon 15, 129–134. was provided by Pedro Viegas and Tim Ziegler. Donna CÚNEO, R., RAMEZANI, J., SCASSO, R., POL, D., ESCAPA, I., ZAVATTIERI, and Terry Tuanui, and Celine and Dianne Gregory-Hunt A.M. & BOWRING, S.A., 2013. High-precision U-Pb geochronol- kindly provided support on the Chatham Islands. Type ogy and a new chronostratigraphy for the Cañadón Asfalto Basin, material was supplied by S. Costanza, Harvard Univer- Chubut, central Patagonia: implications for terrestrial faunal and floral evolution in Jurassic. Gondwana Research 24, 1267–1275. sity Herbaria. Research supported by a National Geo- DETTMANN, M.E., CLIFFORD, H.T. & PETERS, M., 2012. Emwadea graphic Society grant (9761-15) and ANSTO Neutron microcarpa gen. et sp. nov.—anatomically preserved araucarian grant (P5524) awarded to CM; additional financial sup- seed cones from the Winton Formation (late Albian), western – port from the Paleontological Society and Monash Queensland, Australia. Alcheringa 36, 217 237. DONG, C., SUN, B.-N., WU, J.-Y., DU, B.-X., XU, X.-H. & JIN, P.-H., University. CM would like to acknowledge the Moriori 2014. Structure and affinities of Athrotaxites yumenensis sp. nov. people as the original landowners of Rangiaotea (Pitt (Cupressaceae) from the Lower Cretaceous of northwestern Island), and special thanks go to Mr Maui Solomon China. Cretaceous Research 47,25–38. DUNKER,W.,1846. Monographie der Norddeutschen Wealdenbildung. Downloaded by [Naturhistoriska Riksmuseum] at 02:26 05 January 2018 (Executive Chairman, Hokotehi Moriori Trust) for his Oehme and Müller, Braunschweig, 86 pp. assistance with the Moriori language. ESCAPA, I., CÚNEO,R.&AXSMITH, B., 2008. A new genus of the Cupressaceae (sensu lato) from the Jurassic of Patagonia: Implica- tions for conifer megasporangiate cone homologies. Review of Disclosure statement Palaeobotany and Palynology 151,110–122. fl ESCAPA, I.H., GANDOLFO, M.A., CREPET, W.L. & NIXON, K.C., 2016.A No potential con ict of interest was reported by the authors. new species of Athrotaxites (Athrotaxoideae, Cupressaceae) from the Upper Cretaceous Raritan Formation, New Jersey, USA. Bot- any 94, 831–845. Funding ETTINGSHAUSEN, C.F.VON, 1887. Beiträge zur Kenntniss der fossilen flora Neuseelands. Denkschriften der Kaiserlichen Akademie der This work was supported by Australian Nuclear Science and Technol- Wissenschaften, Wien 53, 143–194. ogy Organisation [grant number P5524] and National Geographic FARJON, A., 2010. A Handbook of the World’s Conifers. Brill Aca- Society [grant number 9761-15] demic Publishers, Leiden, Netherlands, 1150 pp. FARJON,A.&ORTIZ GARCIA, S., 2003. Cone and ovule development in Cunninghamia and Taiwania (Cupressaceae sensu lato) and its References significance for conifer evolution. 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