treatments-xml/data/03/9C/64/039C6431083CFFD5A50892A0D4D1FE4D.xml

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<mods:title id="0A849878C3E827C0B1A225884FEA4E9F">The Early Middle Eocene Wagon Bed Carpoflora Of Central Wyoming, U. S. A.</mods:title>
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<mods:namePart id="AEE859853935BF86883E5EF2E552CAD4">Tiffney, Bruce H.</mods:namePart>
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Genus 
<taxonomicName id="4C35AEA4083CFFD7A5F892A0D41EFF00" ID-CoL="3HXZ" authority="NUTT." authorityName="NUTT." box="[419,557,199,223]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A5F892A0D7D5FF00" bold="true" box="[419,486,199,223]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
NUTT.
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<paragraph id="8B8AD527083CFFD7A50B9280D403FF20" blockId="8.[336,560,231,255]" box="[336,560,231,255]" pageId="8" pageNumber="59">
<figureCitation id="130EC9A2083CFFD7A50B9280D7D8FF20" box="[336,491,231,255]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3a–x</figureCitation>
, 
<figureCitation id="130EC9A2083CFFD7A5AD9280D403FF20" box="[502,560,231,255]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">4a–w</figureCitation>
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M a t e r i a l. Ten specimens, four whole, three half nuts and three more fragmentary. Many chalcedony casts. Three specimens exhibiting casts of the seed, 
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permineralized and displaying some internal structure. 
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<specimenCode id="DB937D5C083CFFD7A48093FED758FE6F" box="[219,363,408,433]" collectionCode="PAL" country="Italy" httpUri="http://biocol.org/urn:lsid:biocol.org:col:15292" lsid="urn:lsid:biocol.org:col:15292" name="Herbarium Mediterraneum Panormitanum" pageId="8" pageNumber="59" type="Herbarium">PAL 772346</specimenCode>
, 
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, 
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, 
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, 
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, 
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, 
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, 
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, 
<specimenCode id="DB937D5C083CFFD7A5C293DED7DFFE0E" box="[409,492,441,465]" collectionCode="USNM" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34871" lsid="urn:lsid:biocol.org:col:34871" name="Smithsonian Institution, National Museum of Natural History" pageId="8" pageNumber="59" type="Museum">772355</specimenCode>
, 
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.
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<paragraph id="8B8AD527083CFFD7A4FD938ED7A7F971" blockId="8.[128,768,488,1710]" pageId="8" pageNumber="59">
D e s c r i p t i o n. Nuts ovoid in side view, almost circular in cross section, but with a sense of a four angled symmetry (e.g., 
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), unilocular, with a single seed. Of the eight reasonably complete full or half nuts the average height is 
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(range 
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), the diameter parallel to plane of dehiscence average 
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(range 
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), and the diameter perpendicular to plane of dehiscence average 
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(range 
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). Surface smooth, often marked by eight very fine longitudinal grooves (Textfig. 4b, c, u). Base generally rounded, although in some cases with a slight protrusion. Lateral walls 
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thick, possibly thickening towards the base. Primary septum (perpendicular to plane of dehiscence) 2 to as much as 
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wide at base, tapering to 
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at apex, and extending ~3/4 of the distance to the top of the locule (
<figureCitation id="130EC9A2083CFFD7A58F91CAD4E0FC1A" box="[468,723,941,966]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3e, m, n, s, t, x</figureCitation>
, 
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, h, p, q, v, w
</figureCitation>
). A pair of placental vascular bundles arises at the base of the primary septum, but these bundles soon diverge (e.g., 
<figureCitation id="130EC9A2083CFFD7A4E59669D75CFBF8" box="[190,367,1038,1063]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-fig. 3l, n, v</figureCitation>
), each arching close to the periphery of the primary septum, rather than running parallel to the nut axis. Secondary septum present near base of the nut, and, along with the primary septum, dividing the base of the locule into four lobes (
<figureCitation id="130EC9A2083CFFD7A53796F7D434FB77" box="[364,519,1168,1192]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3n, x</figureCitation>
, 
<figureCitation id="130EC9A2083CFFD7A64996F7D403FB77" box="[530,560,1168,1192]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">4k</figureCitation>
); secondary septum shorter and narrower than the primary septum. Interior of locule smooth with no inner ribs. Locule narrower in the plane of the primary septum (
<quantity id="4CCD78C2083CFFD7A5999696D43FFAD6" box="[450,524,1265,1289]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="59" unit="mm" value="10.0">10 mm</quantity>
) than perpendicular to it (
<quantity id="4CCD78C2083CFFD7A4C49775D6C2FAF5" box="[159,241,1298,1322]" metricMagnitude="-2" metricUnit="m" metricValue="1.4" pageId="8" pageNumber="59" unit="mm" value="14.0">14 mm</quantity>
). Two narrow, continuous, lacunae present in the wall of the nut, originating approximately 1/4 of the way from the base of the fruit and extending to near the apex; possibly bifurcated at the base, but coalescing into a single arcuate cavity in each half of the nutshell, enveloping the locule towards the apex (
<figureCitation id="130EC9A2083CFFD7A5CE97D4D464FA14" box="[405,599,1459,1483]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3j, k, m</figureCitation>
, 
<figureCitation id="130EC9A2083CFFD7A63E97D4D4A0FA14" box="[613,659,1459,1483]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">4i, j</figureCitation>
). Cellular structure present in 
<specimenCount id="9D331EAE083CFFD7A53E97B3D7CDFA33" box="[357,510,1492,1516]" pageId="8" pageNumber="59" type="generic">one specimen</specimenCount>
, where a cross-section reveals the core of the nut to be composed of equiaxial cells. In others, the exterior displays radiating structure that might represent radially-elongate mineral fibers (
<figureCitation id="130EC9A2083CFFD7A6679452D4F8F992" box="[572,715,1589,1613]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3i, j</figureCitation>
, 
<figureCitation id="130EC9A2083CFFD7A68E9452D4C0F992" box="[725,755,1589,1613]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">4q</figureCitation>
). Seed elongate, 
<quantity id="4CCD78C2083CFFD7A5719432D742F9B2" box="[298,369,1621,1645]" metricMagnitude="-2" metricUnit="m" metricValue="1.2" pageId="8" pageNumber="59" unit="mm" value="12.0">12 mm</quantity>
wide × 
<quantity id="4CCD78C2083CFFD7A58E9432D42FF9B2" box="[469,540,1621,1645]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="8" pageNumber="59" unit="mm" value="17.0">17 mm</quantity>
high with a convex shoulder region (
<figureCitation id="130EC9A2083CFFD7A56C9411D7D4F951" box="[311,487,1654,1678]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">Text-fig. 4f, h, l</figureCitation>
); seed coat thin (
<figureCitation id="130EC9A2083CFFD7A6FF9411D6A5F971" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">Text-fig. 4j</figureCitation>
) and apparently papery.
</paragraph>
<paragraph id="8B8AD527083CFFD7A4FD94A1D381FE6D" blockId="8.[128,768,1733,1919]" lastBlockId="8.[843,1482,152,1919]" pageId="8" pageNumber="59">
D i s c u s s i o n. The fossils are clearly nuts of the 
<taxonomicName id="4C35AEA4083CFFD7A4DB9481D722F921" box="[128,273,1766,1790]" class="Magnoliopsida" family="Juglandaceae" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="family">Juglandaceae</taxonomicName>
, with a resemblance to both 
<taxonomicName id="4C35AEA4083CFFD7A6019481D49EF921" box="[602,685,1766,1790]" class="Magnoliopsida" family="Juglandaceae" genus="Juglans" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A6019481D49EF921" box="[602,685,1766,1790]" italics="true" pageId="8" pageNumber="59">Juglans</emphasis>
</taxonomicName>
L. and 
<taxonomicName id="4C35AEA4083CFFD7A4DB9560D6F0F8C1" box="[128,195,1799,1822]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A4DB9560D6F0F8C1" box="[128,195,1799,1822]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
. Some specimens appear to be isolated nuts without preservation of the surrounding husk, but in others the nutshell is permineralized and appears dark in reflected light, and is surrounded by a distinct, but less well preserved lighter colored layer that may represent the husk (as seen in 
<figureCitation id="130EC9A2083CFFD7A7FF92DFD2B3FF0E" box="[932,1152,184,209]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-fig. 3i–m, u, v</figureCitation>
). 
<bibRefCitation id="EFA4A8D6083CFFD7A0CC92DFD310FF0F" author="Scott, R. A." box="[1175,1315,184,208]" pageId="8" pageNumber="59" pagination="66 - 97" refId="ref26222" refString="Scott, R. A. (1954): Fossil fruits and seeds from the Eocene Clarno Formation. - Palaeontographica, Abt. B., Palaophytologie, 96: 66 - 97." type="journal article" year="1954">Scott (1954)</bibRefCitation>
, 
<bibRefCitation id="EFA4A8D6083CFFD7A16A92DED3F6FF0F" author="Leroy, J. F." box="[1329,1477,184,209]" pageId="8" pageNumber="59" pagination="1 - 246" refId="ref24548" refString="Leroy, J. F. (1955): Etude sur les Juglandaceae. A la recherche d'une conception morphologique de la fleur femelle et du fruit. - Memoires du Museum National D'Histoire Naturelle, Serie B, Botanique, 6: 1 - 246." type="journal article" year="1955">Leroy (1955)</bibRefCitation>
, 
<bibRefCitation id="EFA4A8D6083CFFD7A71092BED229FF2E" author="Manchester, S. R." box="[843,1050,217,241]" pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">
<collectingRegion id="49F11BC5083CFFD7A71092BED5F4FF2E" box="[843,967,217,241]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
(1987)
</bibRefCitation>
and 
<bibRefCitation id="EFA4A8D6083CFFD7A00C92BED3FAFF2E" author="Manchester, S. R. &amp; McIntosh, W. C." box="[1111,1481,217,241]" pageId="8" pageNumber="59" pagination="7 - 17" refId="ref25401" refString="Manchester, S. R., McIntosh, W. C. (2007): Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. - PaleoBios, 27: 7 - 17." type="journal article" year="2007">
<collectingRegion id="49F11BC5083CFFD7A00C92BED2E0FF2E" box="[1111,1235,217,241]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
and McIntosh (2007)
</bibRefCitation>
provide characters to distinguish these two genera. While the mode of preservation impedes determination of some of these characters (e.g., dehiscence of the husk, complete path of placental bundles within the nut), others can be evaluated. Our discussion below progresses from characters that suggest affinity with 
<taxonomicName id="4C35AEA4083CFFD7A07093FCD25DFE6D" box="[1067,1134,411,434]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A07093FCD25DFE6D" box="[1067,1134,411,434]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
to those that are less decisive.
</paragraph>
<paragraph id="8B8AD527083CFFD7A72A93DCD220F9C2" blockId="8.[843,1482,152,1919]" pageId="8" pageNumber="59">
The smooth surface seen in the fossils is more commonly characteristic of 
<taxonomicName id="4C35AEA4083CFFD7A04893BCD265FE2D" box="[1043,1110,475,498]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A04893BCD265FE2D" box="[1043,1110,475,498]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
than 
<taxonomicName id="4C35AEA4083CFFD7A0F393BCD2C8FE2C" box="[1192,1275,475,499]" class="Magnoliopsida" family="Juglandaceae" genus="Juglans" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A0F393BCD2C8FE2C" box="[1192,1275,475,499]" italics="true" pageId="8" pageNumber="59">Juglans</emphasis>
</taxonomicName>
, although it can occur in some modern species of the latter. There is a hint of a protuberant base in three of the Wagon Bed fossils similar to the condition in 
<taxonomicName id="4C35AEA4083CFFD7A02D905BD3A0FD8B" authority="(Manchester 1987)" baseAuthorityName="Manchester" baseAuthorityYear="1987" box="[1142,1427,572,596]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A02D905BD28AFD8C" box="[1142,1209,572,595]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
(
<bibRefCitation id="EFA4A8D6083CFFD7A092905BD3B8FD8B" author="Manchester, S. R." box="[1225,1419,572,596]" pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">
<collectingRegion id="49F11BC5083CFFD7A092905BD374FD8B" box="[1225,1351,572,596]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
1987
</bibRefCitation>
)
</taxonomicName>
. The position of the pair of placentary bundles, whether running parallel and adjacent to the central axis as in 
<taxonomicName id="4C35AEA4083CFFD7A114901BD391FD4B" box="[1359,1442,636,660]" class="Magnoliopsida" family="Juglandaceae" genus="Juglans" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A114901BD391FD4B" box="[1359,1442,636,660]" italics="true" pageId="8" pageNumber="59">Juglans</emphasis>
</taxonomicName>
, or diverging from one another near the base and following a peripheral course before converging toward the seed apex as in 
<taxonomicName id="4C35AEA4083CFFD7A79590B9D222FD2A" box="[974,1041,734,757]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A79590B9D222FD2A" box="[974,1041,734,757]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
, is an important distinguishing feature (
<bibRefCitation id="EFA4A8D6083CFFD7A7099099D5E4FCCA" author="Leroy, J. F." box="[850,983,765,790]" pageId="8" pageNumber="59" pagination="1 - 246" refId="ref24548" refString="Leroy, J. F. (1955): Etude sur les Juglandaceae. A la recherche d'une conception morphologique de la fleur femelle et du fruit. - Memoires du Museum National D'Histoire Naturelle, Serie B, Botanique, 6: 1 - 246." type="journal article" year="1955">Leroy 1955</bibRefCitation>
), but the configuration is not well preserved in most of these specimens. However, by moving through virtual slices of the two permineralized specimens shown in 
<figureCitation id="130EC9A2083CFFD7A7339139D268FCA9" box="[872,1115,862,886]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-fig. 3j–n and s–x</figureCitation>
, we were able to follow the faint traces of these bundles. Although the bundles are closely adjacent at the base of the nut (
<figureCitation id="130EC9A2083CFFD7A0F591F8D302FC68" box="[1198,1329,927,951]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-fig. 3n</figureCitation>
), longitudinal sections in plane of the primary septum (
<figureCitation id="130EC9A2083CFFD7A15C91D8D3B2FC08" box="[1287,1409,959,983]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-fig. 3l</figureCitation>
) show that they quickly diverge and assume a peripheral course in the manner of 
<taxonomicName id="4C35AEA4083CFFD7A7B69667D203FBC8" box="[1005,1072,1024,1047]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A7B69667D203FBC8" box="[1005,1072,1024,1047]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
. The seed has a rounded shoulder, is axially elongate within the locule, and is distinctly narrower in plane of primary septum and wider perpendicular to it (
<figureCitation id="130EC9A2083CFFD7A7099606D5FEFBA6" box="[850,973,1121,1145]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">Text-fig. 4l</figureCitation>
), all characteristic of 
<taxonomicName id="4C35AEA4083CFFD7A0EC9606D2C9FBA7" box="[1207,1274,1121,1144]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A0EC9606D2C9FBA7" box="[1207,1274,1121,1144]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFA4A8D6083CFFD7A1539606D3F5FBA6" author="Manchester, S. R." box="[1288,1478,1121,1145]" pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">
<collectingRegion id="49F11BC5083CFFD7A1539606D3B5FBA6" box="[1288,1414,1121,1145]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
1987
</bibRefCitation>
, 
<bibRefCitation id="EFA4A8D6083CFFD7A71096E6D29FFB46" author="Manchester, S. R. &amp; McIntosh, W. C." box="[843,1196,1153,1177]" pageId="8" pageNumber="59" pagination="7 - 17" refId="ref25401" refString="Manchester, S. R., McIntosh, W. C. (2007): Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. - PaleoBios, 27: 7 - 17." type="journal article" year="2007">
<collectingRegion id="49F11BC5083CFFD7A71096E6D5F4FB46" box="[843,967,1153,1177]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
and McIntosh 2007
</bibRefCitation>
). The available characters of the Wagon Bed specimens conform more closely to 
<taxonomicName id="4C35AEA4083CFFD7A71096A5D5BDFB06" box="[843,910,1218,1241]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A71096A5D5BDFB06" box="[843,910,1218,1241]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
than to 
<taxonomicName id="4C35AEA4083CFFD7A7BE96A5D20BFB05" box="[997,1080,1218,1242]" class="Magnoliopsida" family="Juglandaceae" genus="Juglans" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A7BE96A5D20BFB05" box="[997,1080,1218,1242]" italics="true" pageId="8" pageNumber="59">Juglans</emphasis>
</taxonomicName>
and are reminiscent of the specimens from Post, 
<collectingRegion id="49F11BC5083CFFD7A7E49685D23CFB25" box="[959,1039,1250,1274]" country="United States of America" name="Oregon" pageId="8" pageNumber="59">Oregon</collectingRegion>
(
<bibRefCitation id="EFA4A8D6083CFFD7A0479685D35CFB25" author="Manchester, S. R. &amp; McIntosh, W. C." box="[1052,1391,1250,1274]" pageId="8" pageNumber="59" pagination="7 - 17" refId="ref25401" refString="Manchester, S. R., McIntosh, W. C. (2007): Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. - PaleoBios, 27: 7 - 17." type="journal article" year="2007">
<collectingRegion id="49F11BC5083CFFD7A0479685D2A9FB25" box="[1052,1178,1250,1274]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
and McIntosh 2007
</bibRefCitation>
) and the European Cenozoic (
<bibRefCitation id="EFA4A8D6083CFFD7A0779765D2AAFAC5" author="Mai, D. H." box="[1068,1177,1282,1306]" pageId="8" pageNumber="59" pagination="339 - 385" refId="ref24872" refString="Mai, D. H. (1981): Der Formenkreis der Vietnam-Nuss (Carya poilanei (Chev.) Leroy in Europa. - Feddes Repertorium, 92: 339 - 385. https: // doi. org / 10.1002 / fedr. 19810920502" type="journal article" year="1981">
<date id="FF8BF3E7083CFFD7A0779765D2AAFAC5" box="[1068,1177,1282,1306]" pageId="8" pageNumber="59" value="1981-05">Mai 1981</date>
</bibRefCitation>
). These represent one of the oldest occurrences of 
<taxonomicName id="4C35AEA4083CFFD7A06A9744D247FAE5" box="[1073,1140,1315,1338]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A06A9744D247FAE5" box="[1073,1140,1315,1338]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
nuts, the next oldest being those from the late Eocene of 
<collectingRegion id="49F11BC5083CFFD7A00A9724D292FA84" box="[1105,1185,1347,1371]" country="United States of America" name="Oregon" pageId="8" pageNumber="59">Oregon</collectingRegion>
(
<bibRefCitation id="EFA4A8D6083CFFD7A0F49724D5B1FAA4" author="Manchester, S. R. &amp; McIntosh, W. C." pageId="8" pageNumber="59" pagination="7 - 17" refId="ref25401" refString="Manchester, S. R., McIntosh, W. C. (2007): Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. - PaleoBios, 27: 7 - 17." type="journal article" year="2007">
<collectingRegion id="49F11BC5083CFFD7A0F49724D31EFA84" box="[1199,1325,1347,1371]" country="United Kingdom" name="Manchester" pageId="8" pageNumber="59">Manchester</collectingRegion>
and McIntosh 2007
</bibRefCitation>
) and Neogene of Europe (
<bibRefCitation id="EFA4A8D6083CFFD7A0F59704D32DFAA4" author="Mai, D. H." box="[1198,1310,1379,1403]" pageId="8" pageNumber="59" pagination="339 - 385" refId="ref24872" refString="Mai, D. H. (1981): Der Formenkreis der Vietnam-Nuss (Carya poilanei (Chev.) Leroy in Europa. - Feddes Repertorium, 92: 339 - 385. https: // doi. org / 10.1002 / fedr. 19810920502" type="journal article" year="1981">
<date id="FF8BF3E7083CFFD7A0F59704D32DFAA4" box="[1198,1310,1379,1403]" pageId="8" pageNumber="59" value="1981-05">Mai 1981</date>
</bibRefCitation>
). 
<taxonomicName id="4C35AEA4083CFFD7A16E9703D34BFAA4" box="[1333,1400,1380,1403]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A16E9703D34BFAA4" box="[1333,1400,1380,1403]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
is also represented by pollen in the Wagon Bed flora (
<bibRefCitation id="EFA4A8D6083CFFD7A13497E4D207FA63" author="Leopold, E. B. &amp; MacGinitie, H. D." pageId="8" pageNumber="59" pagination="147 - 200" refId="ref24402" refString="Leopold, E. B., MacGinitie, H. D. (1972): Development and affinities of Tertiary floras in the Rocky Mountains. - In: Graham, A. (ed.), Floristics and Paleofloristics of Asia and Eastern North America. Elsevier, Amsterdam, pp. 147 - 200." type="book chapter" year="1972">Leopold and MacGinitie 1972</bibRefCitation>
). In the current day, 
<taxonomicName id="4C35AEA4083CFFD7A14897C3D365FA64" box="[1299,1366,1444,1467]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A14897C3D365FA64" box="[1299,1366,1444,1467]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
comprises some 18–24 species of temperate trees, distributed in North and Central America, with a lesser diversity in eastern Asia (
<bibRefCitation id="EFA4A8D6083CFFD7A7099462D23BF9C2" author="Mabberley, D. J." box="[850,1032,1541,1565]" pageId="8" pageNumber="59" refId="ref24766" refString="Mabberley, D. J. (2008): Mabberley's Plant-Book, 3 rd ed. - Cambridge University Press, Cambridge, 1019 pp." type="book" year="2008">Mabberley 2008</bibRefCitation>
).
</paragraph>
<paragraph id="8B8AD527083CFFD7A72A9442D3FAF8A1" blockId="8.[843,1482,152,1919]" pageId="8" pageNumber="59">
Turning to those characters that are less indicative of 
<taxonomicName id="4C35AEA4083CFFD7A7109421D5BDF982" box="[843,910,1606,1629]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A7109421D5BDF982" box="[843,910,1606,1629]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
, the presence of inner ribs perpendicular to the median septum is characteristic of 
<taxonomicName id="4C35AEA4083CFFD7A0329401D3B3F9A1" authority="(Manchester 1987)" baseAuthorityName="Manchester" baseAuthorityYear="1987" box="[1129,1408,1638,1662]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A0329401D29FF9A2" box="[1129,1196,1638,1661]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
(
<bibRefCitation id="EFA4A8D6083CFFD7A0E19401D34BF9A1" author="Manchester, S. R." box="[1210,1400,1638,1662]" pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">Manchester 1987</bibRefCitation>
)
</taxonomicName>
. These are missing in the fossil, although fig. 3, 
<quantity id="4CCD78C2083CFFD7A14C94E1D377F941" box="[1303,1348,1670,1694]" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="8" pageNumber="59" unit="in" value="1.0">1 in</quantity>
<bibRefCitation id="EFA4A8D6083CFFD7A11694E1D5A1F961" author="Manchester, S. R." pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">Manchester (1987)</bibRefCitation>
suggests these may be very subdued in some cases. This is also true of some living species, e.g., in extant 
<taxonomicName id="4C35AEA4083CFFD7A1DC94A0D228F921" authority="(A. CHEV.)" baseAuthorityName="A. CHEV." class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="species" species="poilanei">
<emphasis id="B9410935083CFFD7A1DC94A0D591F920" italics="true" pageId="8" pageNumber="59">Carya poilanei</emphasis>
(A.CHEV.)
</taxonomicName>
J.-F.LEROY of 
<collectingCountry id="F32295B7083CFFD7A0979481D315F921" box="[1228,1318,1766,1790]" name="Vietnam" pageId="8" pageNumber="59">Vietnam</collectingCountry>
and Southern 
<collectingCountry id="F32295B7083CFFD7A7109561D5B8F8C1" box="[843,907,1798,1822]" name="China" pageId="8" pageNumber="59">China</collectingCountry>
(
<bibRefCitation id="EFA4A8D6083CFFD7A7C69561D23FF8C1" author="Mai, D. H." box="[925,1036,1798,1822]" pageId="8" pageNumber="59" pagination="339 - 385" refId="ref24872" refString="Mai, D. H. (1981): Der Formenkreis der Vietnam-Nuss (Carya poilanei (Chev.) Leroy in Europa. - Feddes Repertorium, 92: 339 - 385. https: // doi. org / 10.1002 / fedr. 19810920502" type="journal article" year="1981">
<date id="FF8BF3E7083CFFD7A7C69561D23FF8C1" box="[925,1036,1798,1822]" pageId="8" pageNumber="59" value="1981-05">Mai 1981</date>
</bibRefCitation>
, 
<bibRefCitation id="EFA4A8D6083CFFD7A0419561D2D5F8C1" author="Zhang, W. - P. &amp; Bai, W. - N. &amp; Zhang, D. - Y." box="[1050,1254,1798,1822]" pageId="8" pageNumber="59" pagination="73 - 92" refId="ref27692" refString="Zhang, W. - P., Bai, W. - N., Zhang, D. - Y. (2022): The rediscovery of Carya poilanei (Juglandaceae) after 80 years reveals a new record from China. - Phytokeys, 188: 73 - 92. https: // doi. org / 10.3897 / phytokeys. 188.77242" type="journal article" year="2022">Zhang et al. 2022</bibRefCitation>
). The nuts of 
<taxonomicName id="4C35AEA4083CFFD7A1DC9561D3F9F8C2" box="[1415,1482,1798,1821]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A1DC9561D3F9F8C2" box="[1415,1482,1798,1821]" italics="true" pageId="8" pageNumber="59">Carya</emphasis>
</taxonomicName>
possess a four-angled amb (
<bibRefCitation id="EFA4A8D6083CFFD7A0269541D30EF8E1" author="Manchester, S. R." box="[1149,1341,1830,1854]" pageId="8" pageNumber="59" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">Manchester 1987</bibRefCitation>
), a character only mildly expressed in two of the ten fossils presented here (
<figureCitation id="130EC9A2083CFFD7A7D39500D238F8A0" box="[904,1035,1895,1919]" captionStart="Text-fig" captionStartId="9.[125,206,1766,1787]" captionTargetBox="[125,1440,159,1751]" captionTargetId="figure-4@9.[123,1451,-46,1987]" captionTargetPageId="9" captionText="Text-fig. 3. Juglandaceae. Carya (a–x). Scale bars = 1cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT" figureDoi="http://doi.org/10.5281/zenodo.7167725" httpUri="https://zenodo.org/record/7167725/files/figure.png" pageId="8" pageNumber="59">Text-figs 3c</figureCitation>
, 
<figureCitation id="130EC9A2083CFFD7A0429500D205F8A0" box="[1049,1078,1895,1919]" captionStart="Text-fig" captionStartId="11.[125,203,1898,1919]" captionTargetBox="[125,1439,155,1882]" captionTargetId="figure-6@11.[89,1454,35,1935]" captionTargetPageId="11" captionText="Text-fig. 4. Juglandaceae Carya (a–w). Scale bars = 1cm. a–d: USNM PAL 772352, reflected light, palladium coated. a: Obliquelateral view of nut, apex up. b: Basal view with damage to left and clear depiction of meridional grooves. c, d: Two lateral views oriented about 130° from each other and avoiding the area of damage; the meridional grooves clear in (c). e–l: USNM PAL 772350. e: Intact nut, lateral view, apex up, reflected light. f: One half of split nut revealing in situ chalcedony locule cast, reflected light. g–k: Virtual sections from micro-CT data. g: Longitudinal section parallel to the exposed face in (f). h: Longitudinal section at 90° from (g). i: Transverse section in apical 1/3 showing locule bracketed by C-shaped lacunae (arrows). j: Equatorial transverse section showing two lobes of the locule separated by primary septum, lacuna evident below as white line. k: Transverse section near base" figureDoi="http://doi.org/10.5281/zenodo.7167727" httpUri="https://zenodo.org/record/7167727/files/figure.png" pageId="8" pageNumber="59">4b</figureCitation>
), although many are casts with some
</paragraph>
<paragraph id="8B8AD527083CFFD7A4DA95A1D2CEF7EC" blockId="8.[129,1482,1990,2099]" pageId="8" pageNumber="59">
<emphasis id="B9410935083CFFD7A4DA95A1D2CEF7EC" bold="true" pageId="8" pageNumber="59">
<taxonomicName id="4C35AEA4083CFFD7A4DA95A1D6D9F804" authorityName="S.R.Manchester" authorityYear="1994" box="[129,234,1990,2011]" class="Magnoliopsida" family="Betulaceae" genus="Coryloides" higherTaxonomySource="GBIF" kingdom="Plantae" order="Fagales" pageId="8" pageNumber="59" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083CFFD7A4DA95A1D6D9F804" bold="true" box="[129,234,1990,2011]" italics="true" pageId="8" pageNumber="59">Coryloides</emphasis>
</taxonomicName>
locule cast, USNM PAL 772345, scale bar = 
<quantity id="4CCD78C2083CFFD7A6E295A1D4DBF804" box="[697,744,1990,2011]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="59" unit="cm" value="1.0">1 cm</quantity>
. o: Lateral view illustrating a disk-like structure at top surmounted by a central point, and the rumpled appearance of the surface, suggesting a possibly flexible texture. p: Lateral view rotated 90°, disk up, the separation between the pointed disk and the rest of the fossil distinct (arrows). q: Lateral view rotated 180° from (p). r: Apical view with circular disk and central point. p–q: Micro-CT scan surface rendering. r: Reflected light, palladium coated.
</emphasis>
</paragraph>
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<paragraph id="8B8AD527083DFFD6A4269481D393F7F7" blockId="9.[125,1441,1766,2088]" pageId="9" pageNumber="60">
Text-fig. 3. 
<taxonomicName id="4C35AEA4083DFFD6A4B59481D748F924" box="[238,379,1766,1787]" class="Magnoliopsida" family="Juglandaceae" kingdom="Plantae" order="Fagales" pageId="9" pageNumber="60" phylum="Tracheophyta" rank="family">Juglandaceae</taxonomicName>
. 
<taxonomicName id="4C35AEA4083DFFD6A5DF9480D7F3F923" box="[388,448,1767,1788]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="9" pageNumber="60" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083DFFD6A5DF9480D7F3F923" bold="true" box="[388,448,1767,1788]" italics="true" pageId="9" pageNumber="60">Carya</emphasis>
</taxonomicName>
(a–x). Scale bars = 1 cm. a–e: USNM PAL 772346. Micro-CT scan surface rendering. a, b: Lateral, c: apical, d: basal views. e: Virtual equatorial transverse section. f–n: USNM PAL 772347. f: Lateral view, reflected light, showing path of saw cut for transverse section of (i). g: Basal view, reflected light. h: Apical view, micro-CT surface rendering. i: Physical transverse section displaying locule and cellular preservation of parts of wall. j–n: Virtual sections from micro-CT scan data. j: Transverse section at apical 1/3 of nut. Note narrow lacunae (arrows). k: Longitudinal section parallel to primary septum, traversing one of the cotyledon lobes and showing secondary septum at base. l: Longitudinal section in plane at right angles to (k) in plane of primary septum, showing divergent placental bundles arising from base of nut (arrows). m: Equatorial transverse section showing two lobes of locule separated by primary septum. n: Transverse section near base of nut showing primary and secondary septa, creating four basal lobes of locule; note diverging placental bundles (arrows). o–x: USNM PAL 772351. o: Lateral view of broken nut with exposed locule cast, reflected light. p: Same orientation of nut, micro-CT surface rendering. q: Same specimen lateral view, rotated 90° from (p), micro-CT surface rendering. r: Apical view, reflected light. s–x: Virtual sections from micro-CT
</paragraph>
</caption>
<paragraph id="8B8AD527083EFFD5A4DA92F0D4D1FE4D" blockId="10.[129,768,151,402]" pageId="10" pageNumber="61">
distortion of the exterior surface. Nuts of 
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<emphasis id="B9410935083EFFD5A61992F0D4A6FF70" box="[578,661,151,175]" italics="true" pageId="10" pageNumber="61">Juglans</emphasis>
</taxonomicName>
generally possess prominent multilobed lacunae in their walls; lacunae are lacking in many species of 
<taxonomicName id="4C35AEA4083EFFD5A5B292BED41FFF2F" box="[489,556,217,240]" class="Magnoliopsida" family="Juglandaceae" genus="Carya" kingdom="Plantae" order="Fagales" pageId="10" pageNumber="61" phylum="Tracheophyta" rank="genus">
<emphasis id="B9410935083EFFD5A5B292BED41FFF2F" box="[489,556,217,240]" italics="true" pageId="10" pageNumber="61">Carya</emphasis>
</taxonomicName>
except in sections 
<emphasis id="B9410935083EFFD5A4DA929ED6DAFECF" box="[129,233,249,272]" italics="true" pageId="10" pageNumber="61">Apocarya</emphasis>
and 
<emphasis id="B9410935083EFFD5A544929ED7B8FECF" box="[287,395,249,272]" italics="true" pageId="10" pageNumber="61">Sinocarya</emphasis>
(
<bibRefCitation id="EFA4A8D6083EFFD5A5C1929FD464FECE" author="Manchester, S. R." box="[410,599,248,273]" pageId="10" pageNumber="61" pagination="1 - 137" refId="ref24948" refString="Manchester, S. R. (1987): The fossil history of the Juglandaceae. - Monographs in Systematic Botany, Missouri Botanical Garden, 21: 1 - 137. https: // doi. org / 10.5962 / bhl. title. 154222" type="journal article" year="1987">
<collectingRegion id="49F11BC5083EFFD5A5C1929FD42BFECF" box="[410,536,248,272]" country="United Kingdom" name="Manchester" pageId="10" pageNumber="61">Manchester</collectingRegion>
1987
</bibRefCitation>
) which possess simple lacunae. Narrow lacunae are readily observed in the permineralized Wagon Bed specimens (e.g., Text figs 3e, j–m, 4i, j), and are present in specimens of 
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<emphasis id="B9410935083EFFD5A603933DD4A8FEAE" box="[600,667,346,369]" italics="true" pageId="10" pageNumber="61">Carya</emphasis>
</taxonomicName>
from the late Eocene of 
<collectingRegion id="49F11BC5083EFFD5A57B931DD743FE4D" box="[288,368,378,402]" country="United States of America" name="Oregon" pageId="10" pageNumber="61">Oregon</collectingRegion>
(
<bibRefCitation id="EFA4A8D6083EFFD5A526931DD4E5FE4D" author="Manchester, S. R. &amp; McIntosh, W. C." box="[381,726,378,402]" pageId="10" pageNumber="61" pagination="7 - 17" refId="ref25401" refString="Manchester, S. R., McIntosh, W. C. (2007): Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. - PaleoBios, 27: 7 - 17." type="journal article" year="2007">
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and McIntosh 2007
</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>