<documentid="108C8B05BB953A0BEE515144B75CC6B2"ID-DOI="10.5070/P9401462457"ID-ISSN="0031-0298"ID-Zenodo-Dep="10913330"IM.bibliography_approvedBy="carolina"IM.illustrations_approvedBy="carolina"IM.materialsCitations_approvedBy="carolina"IM.metadata_approvedBy="felipe"IM.tables_requiresApprovalFor="GgImagineBatch,operationResults"IM.taxonomicNames_approvedBy="carolina"IM.treatments_approvedBy="carolina"checkinTime="1712167746961"checkinUser="felipe"docAuthor="Wheeler, Elisabeth A., Manchester, Steven R. & Baas, Pieter"docDate="2023"docId="038AF505A32A96365546FF03FE629E31"docLanguage="en"docName="PaleoBios.40.14.1-55.pdf"docOrigin="PaleoBios 40 (14)"docSource="http://dx.doi.org/10.5070/p9401462457"docStyle="DocumentStyle:76F55B8832C4952BA7DD61B33E1F0E81.3:PaleoBios.2023-.journal_article"docStyleId="76F55B8832C4952BA7DD61B33E1F0E81"docStyleName="PaleoBios.2023-.journal_article"docStyleVersion="3"docTitle="Carya leroyii Wheeler & Manchester & Baas 2023, SP. NOV."docType="treatment"docVersion="4"lastPageNumber="22"masterDocId="FFB38D7DA33F9621547BFFC0FFD49B4F"masterDocTitle="A late Eocene wood assemblage from the Crooked River Basin, Oregon, USA"masterLastPageNumber="55"masterPageNumber="1"pageNumber="20"updateTime="1728048930804"updateUser="ExternalLinkService"zenodo-license-document="CC-BY-NC-SA-4.0">
<taxonomicNameid="4C233F90A32A96345546FF03FE219B90"authority="Wheeler & Manchester & Baas, 2023"authorityName="Wheeler & Manchester & Baas"authorityYear="2023"box="[317,501,195,223]"class="Magnoliopsida"family="Juglandaceae"genus="Carya"kingdom="Plantae"order="Fagales"pageId="21"pageNumber="49"phylum="Tracheophyta"rank="species"species="leroyii"status="sp. nov.">
<collectionCodeid="ED32DCD6A32A9634567AFF06FDCF9B91"box="[513,539,198,222]"country="Brazil"name="Instituto de Botânica"pageId="21"pageNumber="20"type="Herbarium">SP</collectionCode>
<figureCitationid="13185896A32A963455F4FF2FFDD69A48"box="[399,514,239,263]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">FIG. 10A–I</figureCitation>
Growth rings present. Wood semi-ring-porous. Vessels solitary and in radial multiples of 2–3. Perforations simple, intervessel pits crowded alternate medium to large; vessel-ray parenchyma pits similar in size and shape to intervessel pits. Non-septate fibers. Axial parenchyma scanty paratracheal and in narrow bands throughout the growth rings. Rays 1–3-seriate, heterocellular with procumbent body cells and 1–3 marginal rows of square-upright cells. Crystals in chambered axial parenchyma strands, some crystal containing cells enlarged.
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.
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. Leroy in recognition of the importance of his studies of extant and fossil
<taxonomicNameid="4C233F90A32A9634540BFCEDFEC39808"authorityName="DC ex Perleb"authorityYear="1818"box="[112,279,813,839]"class="Magnoliopsida"family="Juglandaceae"kingdom="Plantae"order="Fagales"pageId="21"pageNumber="20"phylum="Tracheophyta"rank="family">Juglandaceae</taxonomicName>
<figureCitationid="13185896A32A9634568AFC56FF13989B"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10A–C</figureCitation>
<figureCitationid="13185896A32A96345595FC1DFD8D98B8"box="[494,601,989,1015]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10A</figureCitation>
Vessels solitary and in radial multiples of 2–3, average tangential diameter of earlywood vessels 229 (44) µm, range 136–294 µm; perforations exclusively simple (
<figureCitationid="13185896A32A9634568AFB86FF739FCB"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10H</figureCitation>
<figureCitationid="13185896A32A96345582FBAAFDB79FCB"box="[505,611,1130,1156]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10D</figureCitation>
<figureCitationid="13185896A32A96345558FB70FE599F85"box="[291,397,1200,1226]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10E</figureCitation>
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=70) µm, range 339–565 µm; thin-walled tyloses present (
<figureCitationid="13185896A32A96345547FB36FDC59E5E"box="[316,529,1270,1297]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10B, C, F, G, I</figureCitation>
Axial parenchyma scanty paratracheal, in tangential bands one to two cells wide throughout the growth ring, bands more closely spaced in the latewood than in the earlywood (
<figureCitationid="13185896A32A96345578FA66FE589E8E"box="[259,396,1446,1473]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10A, B</figureCitation>
<figureCitationid="13185896A32A963454CCFA0AFE979EAB"box="[183,323,1482,1508]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10G, H</figureCitation>
<figureCitationid="13185896A32A96345515FA2DFDD09D48"box="[366,516,1517,1543]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10 G–H</figureCitation>
<figureCitationid="13185896A32A963454B9F996FE989D3E"box="[194,332,1622,1649]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10E, F</figureCitation>
); uniseriate rays composed of mostly upright cells, total multiseriate ray height averages 438 (
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<figureCitationid="13185896A32A96345615F8C6FD279C6E"box="[622,755,1798,1825]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="21"pageNumber="20">Fig. 10G–I</figureCitation>
<emphasisid="B9579801A32A963454EBF8E9FDBD9C0B"bold="true"box="[144,617,1833,1860]"pageId="21"pageNumber="20">Comparisons with extant woods—</emphasis>
Affinities with
<taxonomicNameid="4C233F90A32A9634540BF88DFEF69C28"authorityName="DC ex Perleb"authorityYear="1818"box="[112,290,1869,1895]"class="Magnoliopsida"family="Juglandaceae"kingdom="Plantae"order="Fagales"pageId="21"pageNumber="20"phylum="Tracheophyta"rank="family">Juglandaceae</taxonomicName>
are indicated by the combination of semi-ring-porosity, vessels solitary and in short radial multiples, narrow continuous lines of axial parenchyma, medium-sized alternate intervessel pits, vessel-ray pa- renchyma pits similar in size to intervessel pits.
Its combination of features, including absence of scalariform perforations (a characteristic of Engelhardieae Mann., 1978), is consistent with placement in the Juglandoideae Eaton (1836) (Wheeler, et al. 2022). Our sample is missing the pith, so we cannot confirm that it was solid, as expected in
Whether crystals are present and their location are useful features in distinguishing genera and species groups in the Juglandoideae. Crystals are common in this wood so it differs from
). The tropical black walnuts have crystalliferous axial parenchyma strands, but not in enlarged cells; the north temperate black walnuts have crystals in enlarged axial parenchyma cells, but not in long chains (
<bibRefCitationid="EFB239E2A32A963451DEFC65FC5498AD"author="Miller, R. B."pageId="21"pageNumber="20"pagination="368 - 377"refId="ref37737"refString="Miller, R. B. 1976. Wood anatomy and identification of species of Juglans. Botanical Gazette 137: 368 - 377. [https: // www. jstor. org / stable / 2473781]."type="journal article"year="1976">Miller 1976</bibRefCitation>
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which are ring-porous and have thick-walled fibers (
<bibRefCitationid="EFB239E2A32A96345043FB59FB149FFB"author="Stark, E. W."box="[1080,1216,1177,1204]"pageId="21"pageNumber="20"refId="ref39572"refString="Stark, E. W. 1953. Wood anatomy of the Juglandaceae indigenous to the United States. Agricultural Experiment Station Bulletin No. 595. Purdue University, Lafayette, Indiana. 42 pp."type="book"year="1953">Stark 1953</bibRefCitation>
).
<bibRefCitationid="EFB239E2A32A963450ACFB59FC439F99"author="Heimsch, C., Jr. & R. H. Wetmore"pageId="21"pageNumber="20"pagination="651 - 660"refId="ref35572"refString="Heimsch, C., Jr. and R. H. Wetmore. 1939. The significance of wood anatomy in the taxonomy of the Juglandaceae. American Journal of Botany 26 (8): 651 - 660. [https: // doi. org / 10.2307 / 2436805]."type="journal article"year="1939">Heimsch and Wetmore (1939)</bibRefCitation>
noted that crystals in swollen axial parenchyma cells were most common in
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.-
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Based on what he acknowledged as limited material, he suggested the two species had similar anatomy. He also noted that the rays in these two species are more heterocellular than in the American species.
<bibRefCitationid="EFB239E2A32A963451EDFA11FBB49D41"author="Muller-Stoll, W. R. & E. Madel"pageId="21"pageNumber="20"pagination="255 - 295"refId="ref37806"refString="Muller-Stoll, W. R. and E. Madel. 1960. Juglandaceen-Holzer aus dem ungarischen Tertiar des pannonischen Becken. Senckenbergiana Lethaea 41: 255 - 295."type="journal article"year="1960">Müller-Stoll and Mädel (1960)</bibRefCitation>
reported another difference for these two species, i.e., they do not have thick-walled vessel elements. However, neither species has crystals in chambered axial parenchyma similar to
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<bibRefCitationid="EFB239E2A32A963457B5F9BEFAB69DD6"author="Heimsch, C., Jr. & R. H. Wetmore"box="[974,1378,1662,1689]"pageId="21"pageNumber="20"pagination="651 - 660"refId="ref35572"refString="Heimsch, C., Jr. and R. H. Wetmore. 1939. The significance of wood anatomy in the taxonomy of the Juglandaceae. American Journal of Botany 26 (8): 651 - 660. [https: // doi. org / 10.2307 / 2436805]."type="journal article"year="1939">Heimsch and Wetmore (1939)</bibRefCitation>
nor
<bibRefCitationid="EFB239E2A32A963451DDF9BFFC439DF3"author="Leroy, J-F."pageId="21"pageNumber="20"pagination="216 - 220"refId="ref36647"refString="Leroy, J-F. 1953. La structure du bois d'Annamocarya. Notes sur le bois des Noyers et autres Juglandacees. Revue Internationale de Botanique Appliquee et d'Agriculture Tropicale 33: 216 - 220."type="journal article"year="1953">Leroy (1953)</bibRefCitation>
gave details about intervessel pitting. The one sample of
we had access to has minute-small intervessel pitting (3–5 µm,
<bibRefCitationid="EFB239E2A32A963450CBF926FA7B9C4E"author="Wheeler, E. A. & P. Baas & S. R. Manchester"box="[1200,1455,1766,1793]"pageId="21"pageNumber="20"pagination="61 - 86"refId="ref40792"refString="Wheeler, E. A., P. Baas and S. R. Manchester. 2022. Wood anatomy of modern and fossil Fagaceae. International Journal of Plant Sciences 183 (1): 61 - 86. [https: // doi. org / 10.1086 / 717328]."type="journal article"year="2022">Wheeler et al 2022</bibRefCitation>
); the only other juglandaceous wood with minute intervessel pitting is
<taxonomicNameid="4C233F90A32A963457CBF8EBFB279C09"authority="Diels and Hand."authorityName="Diels and Hand."box="[944,1267,1835,1863]"class="Magnoliopsida"family="Juglandaceae"genus="Rhoiptelea"higherTaxonomySource="CoL,GBIF"kingdom="Plantae"order="Fagales"pageId="21"pageNumber="20"phylum="Tracheophyta"rank="genus">
<bibRefCitationid="EFB239E2A32A963457DBF88EFB869C27"author="Withner, C. L."box="[928,1106,1870,1896]"pageId="21"pageNumber="20"pagination="872 - 878"refId="ref41427"refString="Withner, C. L. 1941. Stem anatomy and phylogeny of the Rhoipteleaceae. American Journal of Botany 28 (10): 872 - 878."type="journal article"year="1941">Withner 1941</bibRefCitation>
<figureCitationid="13185896A329963754EBF8BBFEC49CDA"box="[144,272,1915,1941]"captionStart="Figure 10"captionStartId="23.[112,187,193,217]"captionTargetId="figure-22@22.[149,1486,192,1896]"captionTargetPageId="22"captionText="Figure 10. Juglandaceae. Carya leroyii sp. nov., UF 278-84908. A, B. Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS. C. Radial multiple of 4, thin-walled tyloses, marginal axial parenchyma, crystal-containing idioblasts, TS. D. Crowded alternate intervessel pits, RLS. E, Vessel-ray parenchyma pitting of similar size as intervessel pits, RLS. F. Body of ray composed of procumbent cells, RLS. G. Thin-walled tyloses in vessel, rays 2‒3 (-4) seriate, crystals in chambered axial parenchyma cells, TLS. H. Oblique end walls of vessel elements (VE), rays 1‒3 (-4) seriate, axial parenchyma strands with crystals. I. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E."figureDoi="http://doi.org/10.5281/zenodo.13890611"httpUri="https://zenodo.org/record/13890611/files/figure.png"pageId="22"pageNumber="21">
<taxonomicNameid="4C233F90A32896365491FF01FEAE9B96"authorityName="DC ex Perleb"authorityYear="1818"box="[234,378,193,217]"class="Magnoliopsida"family="Juglandaceae"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="family">Juglandaceae</taxonomicName>
.
<taxonomicNameid="4C233F90A328963655F8FF03FDDD9B94"authority="Wheeler & Manchester & Baas, 2023"authorityName="Wheeler & Manchester & Baas"authorityYear="2023"box="[387,521,195,219]"class="Magnoliopsida"family="Juglandaceae"genus="Carya"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="species"species="leroyii"status="sp. nov.">
Semi-ring-porous wood with vessels solitary and in radial multiples, axial parenchyma in narrow bands throughout the growth ring, thin-walled tyloses, TS.
. Crystals in enlarged axial parenchyma cell (*) and in chambered axial parenchyma (AP). Scale bars=500 µm in A; 200 µm in B; 100 µm in C, F, G. H, I; 50 µm in D, E.
in chambered axial parenchyma; its heterocellular rays are consistent with earlier observations of Asian species (
<bibRefCitationid="EFB239E2A32896365400FE34FEEE9941"author="Itoh, T. B. & Pan, P. & Baas, J. & Luo, D. & Li, Y. & Cui, F. & Wang & M. Mertz & Y. Yasumoto"box="[123,314,500,526]"pageId="23"pageNumber="22"refId="ref36047"refString="Itoh, T. B. Pan, P. Baas, J. Luo, D. Li, Y. Cui, F. Wang. M. Mertz and Y. Yasumoto. 2022. Anatomical Atlas and Database of Chinese Woods. Kaiseisha Press. 2679 pp."type="book"year="2022">Itoh et al. 2022</bibRefCitation>
). Unfortunately, we do not have information on the Asian
<taxonomicNameid="4C233F90A3289636550CFDD8FEF4991B"authority=", C. C. Cheng and R. H. Chang (1979)"authorityName="C. C. Cheng and R. H. Chang"authorityYear="1979"class="Magnoliopsida"family="Juglandaceae"genus="Carya"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="species"species="hunanensis">
<taxonomicNameid="4C233F90A32896365534FDFBFF139937"authority="Kuang and A. M. Lu (1979)"authorityName="Kuang and A. M. Lu"authorityYear="1979"class="Magnoliopsida"family="Juglandaceae"genus="Carya"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="species"species="kweichowensis">
. Because of its heterocellular rays and abundant crystals, we suggest that
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<emphasisid="B9579801A328963654EBFD07FD8299AE"bold="true"box="[144,598,711,737]"pageId="23"pageNumber="22">Comparisons with fossil woods—</emphasis>
The two juglandaceous woods of the nearby middle Eocene Nut Beds differ:
<taxonomicNameid="4C233F90A328963654BAFCCEFF1C9804"authority="Wheeler and Manchester (2002)"authorityName="Wheeler"authorityYear="2002"class="Magnoliopsida"family="Juglandaceae"genus="Clarnoxylon"higherTaxonomySource="GBIF"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="species"species="blanchardii">
<bibRefCitationid="EFB239E2A32896365591FCCDFF1C9804"author="Wheeler, E. A. & S. R. Manchester"pageId="23"pageNumber="22"pagination="1 - 188"refId="ref41110"refString="Wheeler, E. A. and S. R. Manchester. 2002. Woods of the Eocene Nut Beds flora, Clarno Formation, Oregon, USA. IAWA Journal Supplement 3: 1 - 188."type="journal article"year="2002">Wheeler and Manchester (2002)</bibRefCitation>
</taxonomicName>
has prismatic crystals commonly occurring in enlarged ray parenchyma cells;
<taxonomicNameid="4C233F90A3289636558DFC94FD8298DD"authority="Wheeler and Manchester (2002)"authorityName="Wheeler"authorityYear="2002"class="Magnoliopsida"family="Juglandaceae"genus="Engelhardioxylon"higherTaxonomySource="GBIF"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="species"species="nutbedensis">
<bibRefCitationid="EFB239E2A328963654BAFCB7FD8298DD"author="Wheeler, E. A. & S. R. Manchester"box="[193,598,887,914]"pageId="23"pageNumber="22"pagination="1 - 188"refId="ref41110"refString="Wheeler, E. A. and S. R. Manchester. 2002. Woods of the Eocene Nut Beds flora, Clarno Formation, Oregon, USA. IAWA Journal Supplement 3: 1 - 188."type="journal article"year="2002">Wheeler and Manchester (2002)</bibRefCitation>
</taxonomicName>
has both simple and scalariform perforation plates and prismatic crystals in ray parenchyma.
<taxonomicNameid="4C233F90A3289636550CFC7EFEC998B4"authority="Muller-Stoll and Madel (1960)"authorityName="Muller-Stoll and Madel"authorityYear="1960"class="Magnoliopsida"family="Juglandaceae"genus="Pterocaryoxylon"higherTaxonomySource="GBIF"kingdom="Plantae"order="Fagales"pageId="23"pageNumber="22"phylum="Tracheophyta"rank="genus">
<bibRefCitationid="EFB239E2A32896365635FC7DFEC998B4"author="Muller-Stoll, W. R. & E. Madel"pageId="23"pageNumber="22"pagination="255 - 295"refId="ref37806"refString="Muller-Stoll, W. R. and E. Madel. 1960. Juglandaceen-Holzer aus dem ungarischen Tertiar des pannonischen Becken. Senckenbergiana Lethaea 41: 255 - 295."type="journal article"year="1960">Müller-Stoll and Mädel (1960)</bibRefCitation>
</taxonomicName>
at the nearby Post Hammer locality (
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279) lacks crystals entirely (
<bibRefCitationid="EFB239E2A3289636559BFBC4FF659F0E"author="Wheeler, E. A. & S. R. Manchester"pageId="23"pageNumber="22"pagination="299 - 329"refId="ref41269"refString="Wheeler, E. A. and S. R. Manchester. 2021. A diverse assemblage of late Eocene woods from Oregon, USA. Fossil Imprint 77 (2): 299 - 329. [https: // doi. org / 10.37520 / fi. 2021.022]."type="journal article"year="2021">
possibly two, in the Dietz Hill locality’s carpoflora (
<bibRefCitationid="EFB239E2A328963656F6FB8AFEB69FC7"author="Manchester, S. R. & W. C. McIntosh"pageId="23"pageNumber="22"pagination="7 - 17"refId="ref37371"refString="Manchester, S. R. and W. C. McIntosh. 2007. Late Eocene silicified fruits and seeds from the John Day Formation near Post, Oregon. PaleoBios 27: 7 - 17."type="journal article"year="2007">
The classic treatment of fossil juglandaceous woods is
<bibRefCitationid="EFB239E2A328963654F4FB74FDC49F81"author="Muller-Stoll, W. R. & E. Madel"box="[143,528,1204,1230]"pageId="23"pageNumber="22"pagination="255 - 295"refId="ref37806"refString="Muller-Stoll, W. R. and E. Madel. 1960. Juglandaceen-Holzer aus dem ungarischen Tertiar des pannonischen Becken. Senckenbergiana Lethaea 41: 255 - 295."type="journal article"year="1960">Müller-Stoll and Mädel’s 1960</bibRefCitation>
paper,
<emphasisid="B9579801A3289636560AFB74FF139E5A"italics="true"pageId="23"pageNumber="22">Juglandaceen-Hölzer aus dem ungarischen Tertiär des pannonischen Becken</emphasis>
, which reviewed the anatomy of the family and all fossil woods assigned to the family by that time. None of the woods they described had the combination of features of
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