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Volume 27.1
January–April 2024
Full table of contents
ISSN: 1094-8074, web version;
1935-3952, print version
Recent Research Articles
See all articles in 27.1 January-April 2024
See all articles in 26.3 September-December 2023
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Cassandra L. Knight
Department of Geosciences
Pennsylvania State University
University Park
Pennsylvania 16802 USA
knight.cassi@gmail.com
I graduated in 2012 with an M.S. in Geosciences from Pennsylvania State University, and previously with a B.S. in Geology from Colby College in Maine. My masters research focused on fossil rainforest leaves from Eocene Patagonia, especially the systematics, paleoecology, and paleobiogeography of fossil Laurales. I have also done leaf morphological assessment of the flora from Florissant Fossil Beds National Monument as a Geocorps Intern and have worked doing paleobotanical surveys at Grand Canyon National Park. In addition to Patagonia and the Western USA, I have also done field work in South Africa, New Zealand, and Australia. I now work for the National Park Service, coordinating National Fossil Day.
Peter Wilf
Department of Geosciences
Pennsylvania State University
University Park
Pennsylvania 16802 USA
pwilf@psu.edu
After an eclectic and non-geological undergrad career (B.A. Penn 1985), I spent three years teaching junior high school in New Jersey and then four years freelancing with my guitars in West Philly. I discovered geology and then paleobotany at the early age of 29 and have never looked back. I somehow moved from the street, almost literally, onto the doctoral track in Penn Geology and defended in 1998. Most of my thesis research was done in residence at the Smithsonian, on megafloral and paleoclimatic change across the Paleocene-Eocene boundary in southern Wyoming. During this time and in an ensuing Smithsonian postdoc, I began developing two major subsequent themes of my research: the fossil history of plant-insect associations and the unbelievable riches of Patagonian fossil floras. I spent three terrific years at Michigan, 1999-2002, as a Michigan Fellow and happily joined the Penn State Geosciences faculty in 2002, where I have been developing these and several other wonderful research projects with my students and colleagues all over the world.
APPENDIX 1
Leaf Character Definitions
Unless defined separately below, terminology for leaf scoring (Appendix 3, Appendix 5) followed Ellis et al. (2009).
Midvein thickened at base: qualitative feature. The midvein is considered basally thickened if the midvein in the basal third of the leaf, especially near the insertion point of the petiole, is noticeably thicker than the midvein in the middle and apical portion of the leaf. (Appendix 2, Character 9; Appendix 4, Character 7)
Ratio of midvein w:2° w: the width of the midvein divided by the width of a secondary vein. The width of the secondary vein is measured proximal to the midvein and perpendicular to the secondary vein course (Appendix 2, Character 11; Appendix 4, Character 8).
Basal secondaries acute: This is a measure of acuteness to the midvein of the basalmost pair of secondary veins is, relative to the superjacent pair of secondary veins. The measurement is the difference between the angle of the basalmost pair of secondary veins to the midvein, and the angle of the second pair of secondary veins to the midvein. When the angle measured is equal to zero, the basal secondary veins are not acute, and increasing angle measurements indicate increasing acuteness (Appendix 2, Character 12; Appendix 4, Character 10).
Proximity of secondary loop to margin: Measured as a percentage. The distance from the outermost point of a secondary vein (typically secondary veins in the medial portion of the blade are measured) divided by the total distance between the midvein and the margin, then multiplied by 100. Both lengths are measured along the same line, which is perpendicular to the midvein (Appendix 2, Character 13; Appendix 4, Character 12).
Secondary loop shape: Qualitative feature. The secondary loop shape is scored 'even' if the superjacent and subjacent secondary veins in a secondary loop maintain a nearly constant distance from each other, until they loop near the blade margin. The secondary loop shape is scored 'cone' if the loop width decreases noticeably from the midvein to the blade margin (Appendix 2, Character 14; Appendix 4, Character 13).
Ratio of 2° loop h:w: The maximum height of a secondary loop (measured from the intersection of a secondary with the midvein, to the apex of the secondary loop) divided by the maximum intercostal width of the secondary loop (measured between the intersection of the superjacent secondary vein and the midvein, to where a perpendicular line intersects the subjacent secondary) (Appendix 2, Character 15; Appendix 4, Character 14).
Tooth size: Measured as a percentage. Tooth size is the distance between the tooth sinus and the tooth apex, divided by the total distance of the tooth apex from the midvein, then multiplied by 100. Both lengths are measured along a single line, which is perpendicular to the midvein. The points at which the tooth apex and sinus occur are projected perpendicularly to this single line, and the length between these projected points is measured to calculate the percentage (Appendix 2, Character 24; Appendix 4, Character 22).
APPENDIX 2
Characters used to score the fossil Atherospermophyllum guinazui comb. nov. and living Atherospermataceae, and explanation of coding. The character numbers given in this Appendix correspond to the first column in the scoring matrix (Appendix 3, below).
Character# |
Character name |
Coding |
1 | laminar length:width ratio | Range |
2 | leaf length | range (mm) |
3 | petiole swollen | 0=no, 1=yes |
4 | base shape convex | 0=no, 1=yes |
5 | apex shape straight | 0=no, 1=yes |
6 | apex shape convex | 0=no, 1=yes |
7 | apex shape acuminate | 0=no, 1=yes |
8 | apex angle obtuse | 0=no, 1=yes |
9 | midvein thickened at base | 0=no, 1=yes |
10 | ratio of midvein w:2° w | Range |
11 | secondary attachment to midvein | 0=excurrent, 1=decurrent |
12 | basal secondaries acute | max. angle (degrees) |
13 | proximity of secondary loop to margin | range (%) |
14 | secondary loop shape | 0=even, 1=cone |
15 | ratio of 2° loop h:w | Number |
16 | number of 2° loops | Number |
17 | intersecondaries present | 0=no, 1=yes |
18 | exterior tertiary loops | 0=weak, 1=strong |
19 | exterior tertiary loop size/spacing | 0=irregular, 1=regular |
20 | margin type | 0=entire, 1=toothed |
21 | teeth per cm | Number |
22 | number of tooth orders | Number |
23 | second order teeth compound | 0=no, 1=yes |
24 | tooth size | range (%) |
25 | teeth occur in basal quarter of blade | 0=no, 1=yes |
26 | principal vein branches from outermost tertiary | 0=no, 1=yes |
27 | tooth accessory veins | 0=none, 1=conjunctal |
28 | tooth shape cc/st | 0=no, 1=yes |
29 | tooth shape cc/rt | 0=no, 1=yes |
30 | tooth shape cc/fl | 0=no, 1=yes |
APPENDIX 3
Atherospermataceae Character Matrix
Character data for the fossil Atherospermophyllum guinazui comb. nov. and living Atherospermataceae. Scores for living species are bold where they match or fully capture the fossil species' score or range.
Atherospermophyllum guinazui |
Laurelia novae-zelandiae |
Laureliopsis philippiana |
Laurelia sempervirens |
Atherosperma moschatum |
Nemuaron vieillardii |
Dryadodaphne novoguineensis |
Dryadodaphne trachyphloia |
|
1 |
2.17-3.29 |
1.9-2 |
1.62-3.3 |
2.7-3.5 |
2.8-3 |
1.9-3 |
2.9-3 |
2.6-3.7 |
2 |
50-122 |
35-76 |
52-109 |
53-114 |
62-80 |
41-71 |
74-97 |
48-170 |
3 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
4 |
0,1 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
5 |
1 |
0 |
1 |
1 |
1 |
0 |
0 |
0 |
6 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
7 |
0 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
8 |
0 |
1 |
1 |
0 |
0 |
1 |
0 |
0 |
9 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
10 |
2.75-6.43 |
1.88-4.24 |
1.74-3.04 |
3.04-5.42 |
2.23-3.69 |
3.25-6.21 |
5.35-5.83 |
2.12-5.56 |
11 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
12 |
18 |
5 |
10 |
10 |
6 |
12 |
18 |
12 |
13 |
4-24 |
18-32 |
24-27 |
10-28 |
9-13 |
17-20 |
17-27 |
14-26 |
14 |
0,1 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
15 |
1.47-4.33 |
2.50-4.33 |
3.60-5.40 |
4.00-5.33 |
3.50-5.00 |
2.00-4.50 |
1.86-3.20 |
2.00-2.88 |
16 |
8-16 |
10-13 |
12-15 |
10-16 |
10-14 |
11-15 |
11-17 |
12-16 |
17 |
0,1 |
0 |
1 |
1 |
0 |
0 |
0 |
1 |
18 |
0,1 |
0 |
1 |
1 |
n/a |
n/a |
0 (?) |
1 |
19 |
0,1 |
0 |
1 |
1 |
n/a |
n/a |
0 |
0 |
20 |
1 |
1 |
1 |
1 |
0,1 |
1 |
1 |
1 |
21 |
2-6 |
3-5 |
2-4 |
2-4 |
1-3 |
2-6 |
1-3 |
1-3 |
22 |
1,2 |
2 |
2 |
2 |
2 |
2 |
1 |
1 |
23 |
n/a,1 |
1 |
1 |
1 |
1 |
1 |
n/a |
n/a |
24 |
3.63-11.93 |
6.43-10.34 |
5.27-14.87 |
5.78-7.08 |
7.35-16.13 |
6.72-7.48 |
4.19-6.07 |
2.59-5.73 |
25 |
1 |
1 |
0,1 |
0,1 |
0,1 |
0,1 |
0,1 |
0 |
26 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
1 |
27 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
28 |
0,1 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
29 |
0,1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
30 |
0,1 |
0 |
0 |
0 |
1 |
0 |
0 |
0 |
Number of matches |
7 |
11 |
11 |
9 |
7 |
10 |
9 |
|
Characters scored |
27 |
27 |
27 |
27 |
27 |
27 |
27 |
|
% similarity |
25.93 |
40.74 |
40.74 |
33.33 |
25.93 |
37.04 |
33.33 |
Appendix 3 (continued).
Daphnandra micrantha |
Daphnandra repandula |
Daphnandra apatela |
Doryphora aromatica |
Doryphora sassafras |
||
1 |
2.7-3 |
3-3.7 |
2.66-4.4 |
2.2-3.1 |
2.5-3.92 |
|
2 |
65-156 |
88-159 |
70-123 |
58-139 |
36-124 |
|
3 |
1 |
0 |
0,1 |
1 |
1 |
|
4 |
1 |
1 |
0,1 |
0 |
0 |
|
5 |
1 |
1 |
0,1 |
1 |
0 |
|
6 |
0 |
0 |
0 |
0 |
0,1 |
|
7 |
1 |
1 |
1 |
1 |
1 |
|
8 |
0 |
0 |
0 |
0 |
0 |
|
9 |
0 |
1 |
1 |
0 |
1 |
|
10 |
2-3.32 |
2.19-4.17 |
2.18-4.22 |
2.36-3.94 |
3.12-4.4 |
no matches |
11 |
1 |
1 |
0,1 |
1 |
0,1 |
|
12 |
12 |
10 |
13 |
8 |
19 |
|
13 |
11-24 |
8-17 |
16-25 |
18-27 |
12-18 |
no matches |
14 |
1 |
1 |
1 |
0 |
0 |
no matches |
15 |
2.38-3.40 |
2.41-4.14 |
2.38-4.56 |
1.80-1.88 |
1.63-2.43 |
no matches |
16 |
8-14 |
9-13 |
8-12 |
9-15 |
8-15 |
no matches |
17 |
0 |
0 |
0,1 |
1 |
1 |
|
18 |
0 |
1 |
1 |
0 |
1 |
no matches |
19 |
0 |
0 |
0 |
0 |
0 |
|
20 |
1 |
1 |
1 |
1 |
1 |
|
21 |
2-4 |
2-5 |
2-6 |
1-3 |
1-3 |
|
22 |
2 |
2 |
2 |
1 |
1,2 |
|
23 |
0 |
0 |
1 |
n/a |
n/a,0 |
|
24 |
6.58-8.5 |
2.11-3.4 |
4.65-12.82 |
4.82-5.89 |
5.13-11.99 |
no matches |
25 |
1 |
1 |
1 |
0,1 |
0,1 |
|
26 |
1 |
1 |
1 |
1 |
1 |
|
27 |
1 |
1 |
1 |
1 |
1 |
|
28 |
1 |
1 |
0,1 |
0 |
1 |
|
29 |
1 |
1 |
1 |
1 |
1 |
no matches |
30 |
0 |
0 |
0 |
0 |
0 |
no matches |
9 |
9 |
15 |
9 |
12 |
||
27 |
27 |
27 |
27 |
27 |
||
33.33 |
33.33 |
55.56 |
33.33 |
44.44 |
APPENDIX 4
Characters used to score the fossil Monimiophyllum callidentatum sp. nov. and living Atherospermataceae, and explanation of coding. The character numbers given in this Appendix correspond to the first column in the scoring matrix (Appendix 5, below).
|
Character |
Coding |
1 |
laminar L:W ratio |
number |
2 |
leaf L |
range (mm) |
3 |
petiole swollen |
0=no, 1=yes |
4 |
base shape convex |
0=no, 1=yes |
5 |
base angle acute |
0=no, 1=yes |
6 |
apex angle acute |
0=no, 1=yes |
7 |
midvein thickened at base |
0=no, 1=yes |
8 |
ratio of midvein w:2° w |
range (%) |
9 |
secondary attachment to midvein |
0=excurrent, 1=decurrent |
10 |
basal secondaries acute |
max angle (degrees) |
11 |
secondary loop strength |
0=weak, 1=strong |
12 |
proximity of secondary loop to margin |
range (%) |
13 |
secondary loop shape |
0=even, 1=cone |
14 |
ratio of 2° loop h:w |
number |
15 |
number of 2° loops |
number |
16 |
intersecondaries present |
0=no, 1=yes |
17 |
exterior tertiary loops |
0=weak, 1=strong |
18 |
exterior tertiary loop size/spacing |
0=irregular, 1=regular |
19 |
margin type |
0=entire, 1=toothed |
20 |
teeth per cm |
number |
21 |
number of tooth orders |
number |
22 |
tooth size |
range (%) |
23 |
teeth occur in basal quarter of blade |
0=no, 1=yes |
24 |
tooth accessory veins |
0=none, 1=conjunctal |
25 |
tooth shape cc/fl |
0=no, 1=yes |
APPENDIX 5
Monimiaceae Character Matrix
Character data for living Monimiaceae and the single fossil of Monimiophyllum callidentatum, sp. nov. Scores for living species are bold where they match or fully capture the fossil species score or range.
Monimiophyllum callidentatum |
Kibara coriacea |
Wilkiea rigidifolia |
Kibara moluccana |
Wilkiea angustifolia |
Kairoa suberosa |
Wilkiea hugeliana |
Kibara macrophylla |
Wilkiea austroqueenslandica |
|
1 |
4.56 |
2.31-2.42 |
2.88-3.4 |
2.53-3.17 |
2.25-4.12 |
2.86-3.28 |
2.98-3.82 |
2.83-3 |
2.97-3.43 |
2 |
73 |
155-226 |
161-221 |
205-323 |
70-142 |
177-280 |
49-119 |
63-201 |
86-137 |
3 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
4 |
1 |
0 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
5 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
6 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
7 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
8 |
2.32-3.04 |
1.90-2.20 |
2.29-3.76 |
2.44-3.46 |
2.75-3.83 |
2.88-3.90 |
2.20-3.00 |
1.85-3.62 |
2.16-4.45 |
9 |
0 |
1 |
0 |
1 |
1 |
1 |
0,1 |
1 |
0,1 |
10 |
25-30 |
11 |
23 |
21 |
25 |
27 |
45 |
22 |
10 |
11 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
12 |
8-13 |
19-24 |
7-14 |
16-18 |
15-21 |
8-13 |
16-33 |
11-20 |
16 |
13 |
0 |
1 |
0 |
0 |
1 |
0 |
0 |
0 |
1 |
14 |
1.43-1.6 |
1.64-2.89 |
1.7-2.06 |
1.25-1.32 |
1.14-1.23 |
1.32-1.43 |
1.07-1.89 |
1.33-1.59 |
1.55-2 |
15 |
19 |
9-11 |
15 |
14 |
9-12 |
15-24 |
9-14 |
12-14 |
14-20 |
16 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
1 |
17 |
0 |
0 |
1 |
1 |
1 |
0 |
0,1 |
1 |
1 |
18 |
0 |
0 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
19 |
1 |
0,1 |
0,1 |
0 |
1 |
1 |
1 |
0,1 |
1 |
20 |
5-9 |
1-2 |
2-3 |
n/a |
1-2 |
1-2 |
2-5 |
1-2 |
1-2 |
21 |
1 |
2 |
1 |
n/a |
1 |
1 |
1,2 |
2 |
2 |
22 |
5.03-7.14 |
1.64-2.17 |
1.98-3.41 |
n/a |
2.13-5.38 |
4.16-6.92 |
3.31-8.35 |
3.11-5.98 |
3.58-6.86 |
23 |
1 |
0,1 |
0,1 |
n/a |
1 |
1 |
0,1 |
1 |
0 |
24 |
0 |
1 |
0 |
n/a |
1 |
0 |
1 |
1 |
1 |
25 |
1 |
1 |
1 |
n/a |
1 |
0 |
1 |
0 |
1 |
Number of matches |
10 |
14 |
6 |
11 |
17 |
19 |
13 |
11 |
|
Characters scored |
25 |
25 |
19 |
25 |
25 |
25 |
25 |
25 |
|
% similarity |
40.00 |
56.00 |
31.58 |
44.00 |
68.00 |
76.00 |
52.00 |
44 |
Appendix 5 (continued).
Wilkiea smithii |
Wilkiea sp. "Palmerston" |
Mollinedia schottiana |
Mollinedia widgrenii |
Grazielanthus arkeocarpus |
Mollinedia ovata |
Macropeplus ligustrinus |
Macrotorus utriculatus |
Hennecartia omphalandra |
|
1 |
2.26-3.36 |
2.71-2.94 |
2.13-2.71 |
1.36-3.5 |
2.29-2.86 |
1.85 |
2-2.03 |
2.7-2.78 |
3.62-4.24 |
2 |
61-121 |
103-114 |
38-143 |
53-182 |
20-32 |
135-157 |
36-81 |
178-186 |
76-106 |
3 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
4 |
1 |
0 |
0 |
1 |
0 |
0 |
1 |
0 |
1 |
5 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
6 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
7 |
1 |
1 |
0 |
1 |
0 |
1 |
1 |
0 |
1 |
8 |
3.75-4.40 |
2.14-2.55 |
1.88-2.06 |
2.27-2.52 |
1.50-3.14 |
1.71-2.70 |
2.55-3.44 |
2.37-3.00 |
1.92-3.50 |
9 |
1 |
1 |
0 |
0 |
0 |
1 |
0,1 |
0 |
1 |
10 |
46 |
25 |
17 |
18 |
6 |
9 |
14 |
16 |
11 |
11 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
1 |
12 |
12-16 |
18-23 |
12-17 |
9-13 |
18-23 |
9-14 |
12-18 |
18-21 |
17-25 |
13 |
0 |
0 |
0,1 |
1 |
1 |
1 |
0 |
0,1 |
1 |
14 |
1.18-1.71 |
1.36-1.5 |
1.57-1.75 |
1.76-3.06 |
1.6-2.33 |
1.8-2.15 |
1.60-1.96 |
1.66-1.91 |
1.25-1.44 |
15 |
16-18 |
10-12 |
7-12 |
8-10 |
7-10 |
8 |
9-13 |
8-9 |
16-18 |
16 |
1 |
1 |
0-7 |
0 (?) |
1 |
1 |
0 |
1 |
0 |
17 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
18 |
1 |
1 |
1 |
0 |
1 |
0 |
0 |
1 |
0 |
19 |
0 |
0 |
1 |
0,1 |
1 |
0,1 |
0,1 |
1 |
1 |
20 |
n/a |
n/a |
1-3 |
1-3 |
5-10 |
0-2 |
1-3 |
1 |
2-4 |
21 |
n/a |
n/a |
1 |
1 |
1 |
1 |
1 |
1 |
2 |
22 |
n/a |
n/a |
2.84-6.00 |
3.34-7.73 |
2.36-5.23 |
1.32-4.08 |
4.93-11.31 |
3.29-6.73 |
5.94-8.69 |
23 |
n/a |
n/a |
0,1 |
0 |
0 |
0 |
0 |
0 |
1 |
24 |
n/a |
n/a |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
25 |
n/a |
n/a |
1 |
0 |
1 |
1 |
0 |
0 |
1 |
10 |
7 |
13 |
13 |
10 |
7 |
14 |
8 |
13 |
|
19 |
19 |
25 |
25 |
25 |
25 |
25 |
25 |
25 |
|
52.63 |
36.84 |
52.00 |
52.00 |
40.00 |
28.00 |
56.00 |
32.00 |
52.00 |
Appendix 5 (continued).
Tetrasynandra pubescens |
Steganthera laxiflora |
Steganthera australiana |
Steganthera macooraia |
Steganthera cooperorum |
Steganthera hirsuta |
Matthaea sancta |
Matthaea heterophylla |
Austromatthaea elegans |
|
1 |
2.93-4.21 |
2.2-2.33 |
3.33-3.4 |
2.75-3.07 |
3.23-5.1 |
2.14-2.35 |
2.77-2.96 |
3.75-4.08 |
3.46-5.03 |
2 |
59-167 |
105-112 |
130-136 |
83-110 |
51-126 |
77-310 |
80-119 |
30-53 |
83-176 |
3 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
4 |
1 |
1 |
0 |
0 |
0 |
1 |
0 |
1 |
1 |
5 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
1 |
0 |
6 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
7 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
1 |
8 |
1.56-2.10 |
2.06-2.58 |
2.06-2.65 |
3.33-3.48 |
2.22-3.15 |
1.73-2.14 |
2.00-2.11 |
2.5-4.16 |
1.89-2.59 |
9 |
1 |
1 |
0 |
1 |
1 |
0 |
1 |
0 |
0 |
10 |
14 |
10 |
11 |
9 |
16 |
10 |
12 |
21 |
0 |
11 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
12 |
13-26 |
13-25 |
15-25 |
13-17 |
8-16 |
8-15 |
10-17 |
10-12 |
8-18 |
13 |
1 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
14 |
1.29-1.83 |
1.48-2.56 |
1.18-1.5 |
1.75-1.86 |
1.39-1.57 |
1.17-1.52 |
1.27-1.71 |
1.13-1.4 |
1.27-1.63 |
15 |
8-9 |
6-7 |
11-14 |
12-14 |
8-13 |
10-12 |
9-11 |
12-13 |
14-17 |
16 |
1 |
0 |
1 |
1 |
1 |
0 |
1 |
0 |
1 |
17 |
1 |
1 |
0 |
0 |
1 |
1 |
1 |
1 |
0 |
18 |
1 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
0 |
19 |
1 |
0,1 |
0,1 |
0 |
0 |
0 |
0 |
0 |
1 |
20 |
3-7 |
1-2 |
1-2 |
n/a |
n/a |
n/a |
n/a |
n/a |
2-3 |
21 |
2 |
1 |
1 |
n/a |
n/a |
n/a |
n/a |
n/a |
1,2 |
22 |
3.89-7.45 |
2.42-5.64 |
2.92-8.76 |
n/a |
n/a |
n/a |
n/a |
n/a |
3.68-6.81 |
23 |
1 |
0 |
0 |
n/a |
n/a |
n/a |
n/a |
n/a |
1 |
24 |
1 |
1 |
1 |
n/a |
n/a |
n/a |
n/a |
n/a |
0 |
25 |
1 |
1 |
1 |
n/a |
n/a |
n/a |
n/a |
n/a |
0 |
13 |
10 |
14 |
8 |
9 |
9 |
5 |
8 |
15 |
|
25 |
25 |
25 |
19 |
19 |
19 |
19 |
19 |
25 |
|
52.00 |
40.00 |
56.00 |
42.11 |
47.37 |
47.37 |
26.32 |
42.11 |
60.00 |
Appendix 5 (continued).
Levieria acuminata |
Hedycarya loxocarya |
Hedycarya cupulata |
Hedycarya angustifolia |
Kibaropsis caledonica |
Hedycarya arborea |
Tambourissa purpurea |
Tambourissa elliptica |
Tambourissa ficus |
|
1 |
3.43-3.58 |
2.59-3.34 |
2.47-3.42 |
2.38-3.47 |
2.71-4.52 |
2.53-2.9 |
2.5-2.61 |
1.29-2 |
2.24-2.98 |
2 |
72-136 |
88-147 |
65-116 |
19-118 |
113-233 |
43-84 |
35-99 |
52-145 |
83-176 |
3 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
4 |
0 |
0 |
1 |
1 |
0 |
1 |
0 |
1 |
0 |
5 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
6 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
1 |
7 |
1 |
1 |
1 |
1 |
1 |
1 |
0 |
0 |
0 |
8 |
2.33-3.16 |
1.53-4.52 |
2.79-5.30 |
2.48-4.75 |
4.31-5.85 |
3.69-4.76 |
2.27-4.14 |
3.28-4.83 |
2.37-3.57 |
9 |
1 |
0 |
0 |
0,1 |
0,1 |
0 |
0 |
1 |
1 |
10 |
12 |
20 |
20 |
23 |
0 |
8 |
16 |
14 |
11 |
11 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
0 |
12 |
8-15 |
13-18 |
13-27 |
15-21 |
9-14 |
15-22 |
8-17 |
9-14 |
11-17 |
13 |
1 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
0,1 |
14 |
1.55-2.4 |
1.47-2.06 |
1.36-1.73 |
1.3-1.5 |
1.6-2 |
1.5-2 |
1.47-1.83 |
1.6-2.15 |
1.5-2.67 |
15 |
6-8 |
8-10 |
9-12 |
8-12 |
14-18 |
10-15 |
4-11 |
10-19 |
10-14 |
16 |
0 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
17 |
0 |
0 |
1 |
0 |
1 |
0 |
0 |
0 |
0 |
18 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
19 |
1 |
0,1 |
0,1 |
1 |
0 |
1 |
0 |
0 |
0 |
20 |
1-2 |
1 |
1-2 |
1-4 |
n/a |
1-3 |
n/a |
n/a |
n/a |
21 |
1 |
1 |
1 |
2 |
n/a |
2 |
n/a |
n/a |
n/a |
22 |
3.65-8.57 |
5.29-10.8 |
5.47-10.5 |
2.23-12.1 |
n/a |
6.03-7.86 |
n/a |
n/a |
n/a |
23 |
0 |
0 |
0 |
0 |
n/a |
0 |
n/a |
n/a |
n/a |
24 |
0 |
1 |
0? |
1 |
n/a |
1 |
n/a |
n/a |
n/a |
25 |
0 |
0 |
0 |
1 |
n/a |
0 |
n/a |
n/a |
n/a |
13 |
10 |
14 |
13 |
9 |
13 |
12 |
9 |
6 |
|
25 |
25 |
25 |
25 |
19 |
25 |
19 |
19 |
19 |
|
52.00 |
40.00 |
56.00 |
52.00 |
47.37 |
52.00 |
63.16 |
47.37 |
31.58 |
Appendix 5 (continued).
Tambourissa tau |
Ephippiandra madagascariensis |
Tambourissa leptophylla |
Tambourissa longicarpa |
Decarydendron ranomafanensis |
Xymalos monospora |
Hortonia floribunda |
Palmeria foremanii |
Palmeria scandens |
|
1 |
3.17-3.87 |
1.55-1.88 |
1.5-1.87 |
1.39-3.1 |
1.89-2.16 |
2.12-2.61 |
1.5-3.11 |
2.67-2.93 |
1.81-3.31 |
2 |
165-298 |
34-120 |
60-230 |
71-325 |
36-82 |
55-141 |
39-109 |
44-104 |
102-123 |
3 |
1 |
0 |
0 |
1 |
1 |
0 |
1 |
1 |
0,1 |
4 |
1 |
1 |
1 |
1 |
1 |
0 |
1 |
1 |
1 |
5 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0,1 |
6 |
1 |
0 |
1 |
1 |
1 |
1 |
1 |
1 |
0,1 |
7 |
1 |
0 |
0 |
0 |
0 |
1 |
1 |
1 |
0 |
8 |
2.61-4.49 |
2.50-2.67 |
2.71-4.67 |
1.76-4.16 |
2.40-3.33 |
2.61-4.17 |
1.79-2.46 |
1.78-4.76 |
2.00-2.61 |
9 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
0 |
10 |
13 |
13 |
16 |
13 |
13 |
23 |
14 |
15 |
12 |
11 |
0 |
1 |
0 |
0 |
1 |
1 |
0 |
1 |
0,1 |
12 |
12-18 |
15-30 |
10-21 |
16-26 |
9-15 |
12-17 |
12-18 |
15-18 |
18-25 |
13 |
1 |
1 |
0,1 |
0,1 |
0 |
0 |
0,1 |
0 |
0 |
14 |
1.73-2.24 |
1.7-3.27 |
1.38-2.13 |
1.65-2.65 |
1.5-2.33 |
1.29-1.72 |
1.14-1.76 |
1.29-1.36 |
1.23-2.21 |
15 |
8-15 |
6-8 |
9-13 |
6-15 |
7-13 |
8-12 |
4-6 |
9-14 |
10-14 |
16 |
0 |
0 |
1 |
0 |
1 |
1 |
0 |
0 |
1 |
17 |
0 |
0 |
0 |
1 |
0 |
1 |
0 |
1 |
1 |
18 |
0 |
1 |
0 |
0 |
0 |
0 |
1 |
0 |
0,1 |
19 |
0 |
1 |
0 |
1 |
0 |
0,1 |
0 |
0 |
0 |
20 |
n/a |
1-3 |
n/a |
1-2 |
n/a |
1 |
n/a |
n/a |
n/a |
21 |
n/a |
2 |
n/a |
1 |
n/a |
1 |
n/a |
n/a |
n/a |
22 |
n/a |
6.09-11.32 |
n/a |
8.87-14.42 |
n/a |
4.88-6.52 |
n/a |
n/a |
n/a |
23 |
n/a |
0,1 |
n/a |
0 |
n/a |
0 |
n/a |
n/a |
n/a |
24 |
n/a |
1 |
n/a |
1 |
n/a |
1 |
n/a |
n/a |
n/a |
25 |
n/a |
0 |
n/a |
0 |
n/a |
0 |
n/a |
n/a |
n/a |
9 |
8 |
9 |
12 |
11 |
11 |
11 |
12 |
8 |
|
19 |
25 |
19 |
25 |
19 |
25 |
19 |
19 |
19 |
|
47.37 |
32.00 |
47.37 |
48.00 |
57.89 |
44.00 |
57.89 |
63.16 |
42.11 |
Appendix 5 (continued).
Monimia rotundifolia |
Monimia ovalifolia |
Peumus boldus |
|
1 |
1.35-1.52 |
1.23-1.61 |
1.71-2.32 |
2 |
47-146 |
61-163 |
24-65 |
3 |
1 |
1 |
1 |
4 |
1 |
1 |
1 |
5 |
0 |
0 |
1 |
6 |
0 |
0 |
1 |
7 |
1 |
1 |
1 |
8 |
1.78-2.61 |
2.20-2.57 |
2.06-2.42 |
9 |
0 |
0 |
0 |
10 |
5 |
5 |
22 |
11 |
0 |
0 |
0 |
12 |
15-20 |
13-18 |
12-23 |
13 |
0 |
0 |
0,1 |
14 |
2.04-3.13 |
2.18-4.27 |
1.38-2.2 |
15 |
8-10 |
9-11 |
6-11 |
16 |
0 |
0 |
1 |
17 |
1 |
1 |
0 |
18 |
0 |
0 |
0 |
19 |
0 |
0 |
0 |
20 |
n/a |
n/a |
n/a |
21 |
n/a |
n/a |
n/a |
22 |
n/a |
n/a |
n/a |
23 |
n/a |
n/a |
n/a |
24 |
n/a |
n/a |
n/a |
25 |
n/a |
n/a |
n/a |
8 |
8 |
10 |
|
19 |
19 |
19 |
|
42.11 |
42.11 |
52.63 |
TABLE 1. Macrofossil records of Atherospermataceae and Monimiaceae.
Fossil |
Organ |
Age |
Provenance |
References |
ATHEROSPERMATACEAE |
||||
Laurelites jamesrossii Poole and Francis |
wood |
Campanian |
James Ross Island |
Poole and Francis, 1999 |
Protoatherospermoxylon spp. |
wood |
Late Cretaceous |
Cape Province, S.Africa |
Mädel, 1960; Müller-Stoll and Mädel, 1962 |
Atherospermoxylon bulboradiatum Poole and Gottwald |
wood |
Paleocene |
Seymour Island |
Poole and Gottwald, 2001 |
Atherospermophyllum guinazui (Berry), comb. nov. |
leaves |
Early and middle Eocene |
Patagonia, Argentina |
Berry, 1935, 1938; this paper |
Laurelia-like wood |
wood |
Paleocene-Eocene |
Seymour Island |
Poole and Gottwald, 2001 |
Atherospermataceous wood |
wood |
Late Eocene |
Germany |
Gottwald, 1992 |
Laurelioxylon intermedium Nishida |
wood |
"Tertiary" |
Chile |
Nishida, 1984 |
Atherospermoxylon aegyptiacum (Schenk) R. Kräusel |
wood |
Early Oligocene |
Egypt |
Kräusel, 1939 |
Laurelites doroteaensis Nishida, H. Nishida and Nasa |
wood |
Oligocene-Miocene |
Patagonia, Chile and Argentina |
Nishida et al., 1988; Pujana 2009 |
Laurelia otagoensis Conran, Bannister and D.E. Lee |
leaves with cuticle, fruits |
Early Miocene |
New Zealand |
Conran et al., 2013 |
Atherosperma moschatum Labill. |
leaves |
Pliocene-Pleistocene |
Tasmania |
Hill and MacPhail, 1985 |
MONIMIACEAE |
||||
Hedycaryoxylon hortonioides Mädel |
wood |
Late Cretaceous |
Cape Province, S.Africa |
Mädel, 1960; Müller-Stoll and Mädel, 1962 |
Hedycaryoxylon tambourissoides Poole and Gottwald |
wood |
Campanian |
James Ross Island |
Süss, 1960; Poole and Gottwald, 2001 |
Monimiophyllum antarcticum Zastawniak |
leaves |
?Late Paleocene |
King George Island |
Birkenmajer and Zastawniak, 1989 |
Monimiophyllum callidentatum sp. nov. |
leaf |
Early Eocene |
Patagonia, Argentina |
This paper |
Flacourtioxylon (Monimiaxylon) gifaense |
wood |
Late Eocene |
Libya |
Louvet, 1974 |
Xymaloxylon (Monimiaxylon) zeltenense |
wood |
Early Oligocene |
Libya |
Louvet, 1974 |
Monimiaceous wood |
wood |
Early Oligocene |
Oman |
Privé-Gill et al., 1993 |
TABLE 2. Extant Atherospermataceae species scored.
Species |
Range |
Atherosperma moschatum Labill. |
NSW, Tas, Vic |
Daphnandra apatela Schodde |
NSW, Qld |
Daphnandra micrantha (Tul.) Benth. |
NSW |
Daphnandra repandula (F.Muell) F. Muell. |
NSW, Qld |
Doryphora aromatica (F.M.Bailey) L.S. Sm. |
Qld |
Doryphora sassafras Endl. |
NSW, Qld, Vic |
Dryadodaphne novoguineensis (Perk.) A.C. Sm. |
New Guinea |
Dryadodaphne trachyphloia Schodde |
Qld |
Laurelia novae-zelandiae A. Cunn. |
New Zealand |
Laurelia sempervirens (Ruiz and Pav.) Tul. |
S Chile |
Laureliopsis philippiana (Looser) Schodde |
S Chile and Argentina |
Nemuaron vieillardii (Baill.) Baill. |
New Caledonia |
Abbreviations: NSW- New South Wales, Tas- Tasmania, Vic- Victoria, Qld- Queensland, Australia.
TABLE 3. Extant Monimiaceae species scored.
Species |
Range |
Austromatthaea elegans L.B. Sm. |
Qld |
Decarydendron ranomafanensis Lorence and Razafim. |
Madagascar |
Ephippiandra madagascariensis (Danguy) Lorence |
Madagascar |
Grazielanthus arkeocarpus Peixoto and Per.-Moura |
SE Brazil |
Hedycarya angustifolia A. Cunn. |
Qld, Tas |
Hedycarya arborea J.R. Forst. and G. Forst. |
New Zealand |
Hedycarya cupulata Baill. |
New Caledonia |
Hedycarya loxocarya (Benth.) W.D. Francis |
Qld |
Hennecartia omphalandra Poiss. |
S Brazil, NE Argentina, Paraguay |
Hortonia floribunda Wight ex Arn. |
Sri Lanka |
Kairoa suberosa Philipson |
New Guinea |
Kibara coriacea (Blume) Hook. f. and A. Thomps. |
Malaysia |
Kibara macrophylla (A. Cunn.) Benth. |
NSW |
Kibara moluccana Boerl. ex Perk. |
New Guinea, Moluccas |
Kibaropsis caledonica (Guillaumin) J. Jeremie |
New Caledonia |
Levieria acuminata Perk. |
New Guinea, Qld |
Macropeplus ligustrinus (Tul.) Perk. |
SE Brazil |
Macrotorus utriculatus Perk. |
SE Brazil |
Matthaea sancta Blume |
Malaysia, Indonesia |
Mollinedia ovata Ruiz and Pav. |
Ecuador |
Mollinedia schottiana (Spreng.) Perk. |
Brazil |
Mollinedia widgrenii A. DC. |
Brazil |
Monimia ovalifolia Thouars |
Réunion, Mascarenes |
Monimia rotundifolia Thouars |
Réunion, Mascarenes |
Palmeria foremanii Whiffin |
NSW, Qld |
Palmeria scandens F. Muell. |
NSW, Qld |
Peumus boldus Molina |
Chile |
Steganthera australiana C.T. White |
New Guinea |
Steganthera cooperorum Whiffin |
Qld |
Steganthera hirsuta Perk. |
Qld, New Guinea |
Steganthera laxiflora (Benth.) Whiffin and Foreman |
Qld |
Steganthera macooraia (F.M. Bailey) P.K. Endress |
Qld |
Tambourissa elliptica A. DC. |
Réunion, Mascarenes |
Tambourissa ficus (Tul.) A. DC. |
Mauritius, Mascarenes |
Tambourissa leptophylla (Tul.) A. DC. |
Mayotte, Comoros |
Tambourissa longicarpa Lorence |
Madagascar |
Tambourissa purpurea (Tul.) A. DC. |
Mauritius, Mascarenes |
Tambourissa tau Lorence |
Mauritius, Mascarenes |
Wilkiea angustifolia (F.M. Bailey) Perk. |
Qld |
Wilkiea austroqueenslandica Domin |
NSW, Qld |
Wilkiea hugeliana (Tul.) A. DC. |
NSW, Qld |
Tetrasynandra pubescens Perk. |
Qld |
Wilkiea rigidifolia (A.C. Sm.) Whiffin and Foreman |
Qld, New Guinea |
Wilkiea smithii Whiffin |
Qld |
Wilkiea sp. "Palmerston" |
Qld |
Xymalos monospora (Harv.) Baill. |
Bioko, Cameroon |
Abbreviations: NSW- New South Wales, Tas- Tasmania, Vic- Victoria, Qld- Queensland, Australia.
TABLE 4. Percent character similarity of extant Atherospermataceae to Atherospermophyllum guinazui, comb. nov.
Taxon | %Similarity |
Genus | |
Daphnandra | 63 |
Doryphora | 56 |
Laurelia | 56 |
Dryadodaphne | 48 |
Laureliopsis | 41 |
Atherosperma | 33 |
Nemuaron | 26 |
Species | |
Daphnandra apatela | 56 |
Doryphora sassafras | 44 |
Laureliopsis philippiana | 41 |
Laurelia sempervirens | 41 |
Dryadodaphne novoguineensis | 37 |
Atherosperma moschatum | 33 |
Dryadodaphne trachyphloia | 33 |
Daphnandra micrantha | 33 |
Daphnandra repandula | 33 |
Doryphora aromatica | 33 |
Laurelia novae-zelandiae | 26 |
Nemuaron vieillardii | 22 |
Notes: Generic scores are a compilation of the constituent species scores. All taxa scored have toothed leaf-margins.
TABLE 5. Percent character similarity of extant Monimiaceae to Monimiophyllum callidentatum, sp. nov.
Taxon |
%Similarity |
Genus |
|
Wilkiea^ |
76 |
Mollinedia^ |
76 |
Kibara^ |
68 |
Kairoa* |
68 |
Hedycarya^ |
68 |
Tambourissa^ |
68 |
Palmeria |
68 |
Steganthera^ |
64 |
Austromatthaea* |
60 |
Decarydendron |
58 |
Hortonia |
58 |
Macropeplus^ |
56 |
Peumus |
53 |
Hennecartia* |
52 |
Levieria* |
52 |
Kibaropsis |
47 |
Xymalos^ |
44 |
Monimia |
42 |
Grazielanthus* |
40 |
Macrotorus* |
32 |
Ephippiandra* |
32 |
Matthaea |
26 |
Species |
|
Wilkiea hugeliana* |
76 |
Kairoa suberosa* |
68 |
Tambourissa purpurea |
63 |
Palmeria foremanii |
63 |
Austromatthaea elegans* |
60 |
Decarydendron ranomafanensis |
58 |
Hortonia floribunda |
58 |
Wilkiea rigidifolia* |
56 |
Macropeplus ligustrinus* |
56 |
Steganthera australiana* |
56 |
Hedycarya cupulata* |
56 |
Wilkiea smithii |
53 |
Peumus boldus |
53 |
Kibara macrophylla* |
52 |
Mollinedia schottiana* |
52 |
Mollinedia widgrenii* |
52 |
Hennecartia omphalandra* |
52 |
Tetrasynandra pubescens* |
52 |
Levieria acuminata* |
52 |
Hedycarya angustifolia* |
52 |
Hedycarya arborea* |
52 |
Tambourissa longicarpa* |
48 |
Steganthera cooperorum |
47 |
Steganthera hirsuta |
47 |
Kibaropsis caledonica |
47 |
Tambourissa elliptica |
47 |
Tambourissa tau |
47 |
Tambourissa leptophylla |
47 |
Wilkiea angustifolia* |
44 |
Xymalos monospora* |
44 |
Wilkiea austroqueenslandica |
44 |
Steganthera macooraia |
42 |
Palmeria scandens |
42 |
Monimia rotundifolia |
42 |
Monimia ovalifolia |
42 |
Matthaea heterophylla |
42 |
Kibara coriacea* |
40 |
Grazielanthus arkeocarpus* |
40 |
Steganthera laxiflora* |
40 |
Hedycarya loxocarya* |
40 |
Wilkiea sp. "Palmerston" |
37 |
Kibara moluccana |
32 |
Tambourissa ficus |
32 |
Macrotorus utriculatus* |
32 |
Ephippiandra madagascariensis* |
32 |
Mollinedia ovata* |
28 |
Matthaea sancta |
26 |
Notes: Generic scores are a compilation of the constituent species scores.
*Genus or species with toothed leaves.
^Genus in which some species are toothed.
FIGURE 1. Ranges of extant genera in 1 - Atherospermataceae and 2 - Monimiaceae. Extant ranges are indicated by blue fill, and red stars indicate the early Eocene Laguna del Hunco and middle Eocene Río Pichileufú fossil localities, from which Atherospermophyllum guinazui (Berry) comb. nov. and Monimiophyllum callidentatum sp. nov. are here reported. The living ranges of Laurelia-Laureliopsis in South America are close to but do not include the fossil sites, which are located slightly to their east.
FIGURE 2. Species character scores for Atherospermataceae, including the fossil Atherospermophyllum guinazui, comb. nov., plotted against a molecular phylogeny for the family (redrawn from Renner et al., 2000). Characters as numbered across the top of the plot correspond to Appendix 2. Circles indicate discrete (presence/absence) characters, white = absent, black = present, half-filled = both absent and present in the species. Squares indicate continuous characters, filled when the scored range of the living species includes the range scored for A. guinazui. Based on these scores, percent similarity of each living species to the fossil species A. guinazui is plotted in the bar graph at right.
FIGURE 3. Species character scores for Monimiaceae, including the fossil Monimiophyllum callidentatum, sp. nov., plotted against a molecular phylogeny for the family (redrawn from Renner et al., 2010), including unscored taxa. Summary scores for additional taxa (not analyzed by Renner et al., 2010) shown in Table 5, Appendix 4, and Appendix 5. Characters numbered across the top of the plot correspond to Appendix 3. Circles indicate discrete (presence/absence) characters, white = absent, black = present, half-filled = both absent and present in the species. Squares indicate continuous characters, filled when the character range of the living species includes the range exhibited by M. callidentatum. Based on these scores, percent similarity of each living species to the fossil species M. callidentatum is plotted in the bar graph at right. Black bars = toothed species, white bars = untoothed species.
FIGURE 4. Atherospermophyllum guinazui (Berry), comb. nov. 1 - Lectotype, USNM 40403a, from Río Pichileufú (RP); 2 - camera lucida drawing of USNM 40403a, showing venation and tooth details; 3 - USNM 40403b (syntype); 4 - camera lucida drawing of USNM 40403b. Selected new specimens of A. guinazui: 5 - MPEF-Pb 5639 (Laguna del Hunco, LH); 6 - MPEF-Pb 5650 (LH); 7 - MPEF-Pb 5640 (LH); 8 - BAR 4732 (RP); 9 - MPEF-Pb 5648 (LH); 10 - MPEF-Pb 5636 (LH). All scale bars are 1 cm. Also visible on USNM 40403a is extensive insect hole and margin feeding and skeletonization.
FIGURE 5. Atherospermophyllum guinazui (Berry), comb. nov., venation (1, 3, 5) and tooth (2, 4, 6) details. 1 and 2 - USNM 40403a (lectotype, Río Pichileufú); 3 and 4 - MPEF-Pb 5639 (Laguna del Hunco, LH); and 5 and 6 - MPEF-Pb 5641 (LH). Scale bar is 1 cm for 1, 3, and 5, and 1 mm for 2, 4, and 6.
FIGURE 6. Atherospermophyllum guinazui (Berry), comb. nov., exemplars and visual comparisons to living Atherospermataceae species (Figure 2, Table 4). The left column (1-3) shows fossil specimens (Figure 4), and the four other columns, left to right, each show herbarium specimens of Daphnandra apatela (4-6), Doryphora sassafras (7-9), Laureliopsis philippiana (10-12), and Laurelia sempervirens (13-15). Fossil specimens: 1 – MPEF-Pb 5639 (LH); 2 – USNM 40403a (lectotype, RP); and 3 – MPEF-Pb 5650 (LH). Examples of living Daphnandra apatela: 4 – NSW 67890, C. Moore 167, Richmond River, NSW, Australia; 5 –CANB 128677, R. Schodde 3366, Sarabah Range, Qld., Australia; and 6 – NSW 102541, R. Schodde 5123, Doyle's River State Forest, NSW, Australia. Doryphora sassafras: 7 – NSW 67954, F. M. Bailey, NSW, Australia; 8 – NSW 69329, R. Schodde 3199, Williams River, NSW, Australia; and 9 – CANB 00467700, P. Martensz 222, Mt. Keira, NSW, Australia. Laureliopsis philippiana: 10 – NSW 67915, A. Castellanos, Lago Menéndez, Argentina; 11 – NSW s/n, B.J. Wallace 239/85; and 12 – NSW 67915, A. Castellanos, Lago Menéndez, Argentina. Laurelia sempervirens: 13 – GH s/n, T. Plowman 2618, Isla Tenglo, Chile; 14 – GH s/n, A.L. Cabrera 6026, Laguna Frias, Argentina; and 15 – NSW 618673, J. Allen s/n, cultivated, Mt. Tomah, NSW, Australia. Scale bars are 1 cm.
FIGURE 7. Monimiophyllum callidentatum, sp. nov., holotype, MPEF-Pb 5630 (Laguna del Hunco), part and counterpart (1 and 2), and details of the toothed margin (3) and the base (4). Note the deep midvein impression visible on the counterpart. Scale bar is 1 cm for 1 and 2, and 1 mm for 3 and 4.
FIGURE 8. Monimiophyllum callidentatum, sp. nov., holotype and visual comparisons to living Monimiaceae species (Figure 3, Table 5). The left-most image shows the fossil specimen (Figure 7; part), and all other images show herbarium specimens. 1 – M. callidentatum holotype from Laguna del Hunco, MPEF-Pb 5630; 2 – Wilkiea hugeliana, C.L. Knight 22, Gibbergunya Range Road near Bigbox Road, Border Ranges N.P., NSW, Australia; 3 – Kairoa suberosa, CANB 204095, R. Schodde 5412, Mt. Moiba, Papua, Indonesia; 4 – Austromatthaea elegans, CANB 128486, R. Schodde 3233, Davies Creek Forestry Reserve, Queensland, Australia; 5 – Hedycarya cupulata, NSW s/n, G. McPherson 2426, Thy River Basin, New Caledonia; 6 – Wilkiea rigidifolia, CANB 173624, R. Pullen 7231, Western District, Papua New Guinea; 7 – Macropeplus ligustrinus, P 02005903, R. M. Harley 25112, Minas Gerais, Brazil (downloaded image, used with permission). Scale bars are 1 cm.
Rare leaf fossils of Monimiaceae and Atherospermataceae (Laurales) from Eocene Patagonian rainforests and their biogeographic significance
Plain Language Abstract
This study focuses on two fossil-leaf species of flowering plants found at the Eocene fossil localities Laguna del Hunco (LH, ca. 52.2 Ma) and Río Pichileufú (RP, ca. 47.7 Ma) in Patagonia, Argentina. These sites yield some of the most diverse assemblages of fossil plants known and have been the focus of many paleontological studies. Historically, these paleofloras were thought to include several fossil plant species with affinities to living forests in southern South America. However, many of the taxa have recently been reclassified into genera with modern-day ranges in Australasian rainforests. Thus, the fossil floras may not include many connections to living Patagonian floras and instead support links to trans-Antarctic Gondwanan floras of the warm Eocene.
The two fossil species studied here, based on characteristic leaf morphology, show affinity to the magnoliid order Laurales of the basal angiosperms. Laurales are a diverse order with many genera that today show distinctly Gondwanan, often disjunct Southern Hemisphere distributions, making them of key interest for understanding southern biogeography. However, Laurales are well represented in the fossil record by only one of its families, Lauraceae. Here, we report new fossil evidence of Laurales from the families Atherospermataceae and Monimiaceae. One fossil species of the family Atherospermataceae, Laurelia guinazui, was previously described by Berry (1935) from RP, but new specimens from LH and RP with better preservation prompted our re-evaluation. The second fossil leaf species, of the family Monimiaceae, is new and known from a single specimen only. The purpose of this study is similar to that of other work evaluating certain components of the floras: to identify the fossil species to at least family level, to determine their probable closest living relatives, and to use both the fossil occurrences and the living ranges of relatives to improve understanding of paleobiogeography and ancient ecology.
Our results suggest that the most similar living relatives of Laurelia guinazui, reassigned here to Atherospermophyllum guinazui (Berry) comb. nov. (Atherospermataceae), and the new fossil species, Monimiophyllum callidentatum sp. nov. (Monimiaceae), live only in Australasia today. These records of Atherospermataceae and Monimiaceae are important because they are some of the only fossil occurrences of the two families from South America known to date. The affinities of these two Patagonian fossil species with living Australian taxa further support the recent evidence that the Patagonian fossil floras show strongest connection to living Australasian rainforest floras, and that significant biotic exchange was occurring between South America and Australasia, via Antarctica, before the final separation of South America and Antarctica during the middle-late Eocene. Based on fossil leaf traits and the ecology of the living relatives, our results also reveal a novel understory component of basal angiosperms on the Eocene landscape.
Resumen en Español
Hojas fósiles de Monimiaceae y Atherospermataceae (Laurales) de los bosques húmedos del Eoceno de Patagonia y su significado biogeográfico
Laguna del Hunco (~52.2 Ma) y Río Pichileufú (~ 47.7 Ma) son dos localidades del Eoceno de Patagonia (Argentina) que han proporcionado algunas de las floras fósiles más diversas de los bosques húmedos, dominados por angiospermas, de Gondwana. En este trabajo nos centramos en el estudio de hojas fósiles dentadas correspondientes a las familias Atherospermataceae y Monimiaceae (Laurales), que actualmente muestran una amplia, y a menudo disyunta, distribución en el hemisferio sur y que presentan un particular interés para la biogeografía de Gondwana. Damos a conocer 24 nuevos ejemplares de Laurelia guinazui Berry 1935 (Atherospermataceae), especie de la que proponemos su reasignación a Atherospermophyllum gen. nov. y A. guinazui (Berry) comb. nov. Se creía que esta especie era un elemento sudamericano de las floras del Eoceno, pero hemos descubierto que muestra mayor afinidad con los géneros australianos actuales, estrechamente relacionados, Daphnandra y Doryphora que con Laureliopsis (Sudamérica) y Laurelia (Sudamérica y Nueva Zelanda). La familia Monimiaceae está representada por un único ejemplar, procedente de Laguna del Hunco, y que atribuimos a Monimiophyllum callidentatum sp. nov. Este fósil muestra una gran semejanza con Wilkiea, un género descendiente actual, en clara contradicción con los análisis moleculares que sitúan la divergencia del clado Wilkiea en Australasia hace 16-38 Ma. Por tanto, el linaje Wilkiea podría ser más antiguo y tener una historia biogeográfica más dilatada a lo largo de Gondwana. Nuestros hallazgos mejoran significativamente el escaso registro fósil de Atherospermataceae y Monimiaceae. Las formas fósiles se hallan a una gran distancia actual de sus parientes vivos más cercanos y, especialmente, con géneros similares asociados, lo que aumenta los lazos de las floras eocenas de Patagonia con Australasia y debilita aún más sus señales de supervivencia en los bosques actuales de Sudamérica.
Palabras clave: Patagonia; Eoceno; Laurales; biogeografía; nuevo género; nueva especie
Traducción: Miguel Company
Résumé en Français
De rares fossiles de feuilles de Monimiaceae et Atherospermataceae (Laurales) des forets tropicales éocènes de Patagonie et leur importance biogéographique
Deux site fossiles éocènes de Patagonie, Argentine, Laguna del Hunco (ca. 52,2 Ma) et Río Pichileufú (ca. 47,7 Ma), ont donné parmi les flores fossiles les plus diversifiés connues à ce jour, représentant des forêts tropicales gondwaniennes dominées par les angiospermes. Nous mettons l'accent sur de rares fossiles de feuilles dentées appartenant aux familles Atherospermataceae et Monimiaceae (Laurales), qui montrent actuellement une large distribution au sud, souvent discontinue, et présentent un grand intérêt pour la biogéographie gondwanienne. Pour Laurelia guinazui Berry 1935 (Atherospermataceae), nous présentons 24 nouveaux spécimens et proposons la réaffectation à Atherospermophyllum gen. nov. et A. guinazui (Berry) comb. nov. Cette espèce était considérée comme un élément sud-américain des flores Eocènes, mais nous montrons une plus forte ressemblance avec les genres actuels australiens d'affinité proche Daphnandra et Doryphora plutôt qu'avec Laureliopsis (Amérique du Sud) et Laurelia (Amérique du Sud et Nouvelle Zélande). Les Monimiaceae sont représentées par un seul spécimen fossile de Laguna del Hunco, attribué ici à Monimiophyllum callidentatum sp. nov. Ce fossiles montrent de grandes similarités avec Wilkiea, un genre dérivé actuel d'Australie, en contradiction apparente avec les analyses moléculaires plaçant la divergence du clade Wilkiea en Australasie vers 16-38 Ma. En conséquence, la lignée Wilkiea pourrait être plus ancienne et avoir une histoire biogéographique plus large à l'échelle du Gondwana. Nos découvertes améliorent significativement le registre fossile pauvre pour les Atherospermataceae et les Monimiaceae. Les fossiles ont été trouvés à bonne distance de leur plus proches parents actuels et, remarquablement, avec des genres associés similaires, augmentant les liens entre les flores de l'Eocène de Patagonie et d'Australasie et affaiblissant d'autant leur signal de survie dans les forêts actuelles d'Amérique du Sud.
Mots clés : Patagonie; Eocène; Laurales; biogéographie; nouveau genre; nouvelles espèces
Translator: Olivier Maridet
Deutsche Zusammenfassung
In progress
Translator: Eva Gebauer
Arabic
Translator: Ashraf M.T. Elewa
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Review: The Princeton Field Guide to Mesozoic Sea Reptiles
The Princeton Field Guide to Mesozoic Sea Reptiles
Article number: 26.1.1R
April 2023 -