<document id="82A14169F30ADB07544044D59FDE49F8" ID-DOI="10.11646/phytotaxa.203.2.1" ID-ISSN="1179-3163" ID-Zenodo-Dep="13638671" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1725340430123" checkinUser="felipe" docAuthor="Huang, Ya-Yi, Mori, Scott A. &amp; Kelly, Lawrence M." docDate="2015" docId="03B0445EFFDCFFCEFF19698E571A8695" docLanguage="en" docName="phytotaxa.203.2.1.pdf" docOrigin="Phytotaxa 203 (2)" docSource="http://dx.doi.org/10.11646/phytotaxa.203.2.1" docStyle="DocumentStyle:96748F8F1B6C902996E134952A3A36B9.13:Phytotaxa.2014-.journal_article" docStyleId="96748F8F1B6C902996E134952A3A36B9" docStyleName="Phytotaxa.2014-.journal_article" docStyleVersion="13" docTitle="Lecythis poiteaui O. Berg 1858" docType="treatment" docVersion="2" lastPageNumber="99" masterDocId="FF893C26FFD2FFC0FF916910533D8538" masterDocTitle="Toward a phylogenetic-based Generic Classification of Neotropical Lecythidaceae- I. Status of Bertholletia, Corythophora, Eschweilera and Lecythis" masterLastPageNumber="121" masterPageNumber="85" pageNumber="99" updateTime="1725346518253" updateUser="ExternalLinkService" zenodo-license-figures="UNSPECIFIED" zenodo-license-treatments="UNSPECIFIED">
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<mods:title id="188A3862C007F0C94F117F2C0B790EB6">Toward a phylogenetic-based Generic Classification of Neotropical Lecythidaceae- I. Status of Bertholletia, Corythophora, Eschweilera and Lecythis</mods:title>
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<mods:namePart id="E60E6B8739C7B950456F883873F3D1F8">Huang, Ya-Yi</mods:namePart>
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<paragraph id="8BA6F548FFDCFFCEFF19698E51A98581" blockId="14.[136,660,158,185]" box="[136,660,158,185]" pageId="14" pageNumber="99">
<heading id="D0EE4224FFDCFFCEFF19698E51A98581" box="[136,660,158,185]" fontSize="11" level="2" pageId="14" pageNumber="99" reason="3">
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<taxonomicName id="4C198ECBFFDCFFCEFF19698E52718580" ID-CoL="6P5VT" authorityName="O. Berg" authorityYear="1858" box="[136,332,158,185]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="poiteaui">
<emphasis id="B96D295AFFDCFFCEFF19698E52718580" bold="true" box="[136,332,158,185]" italics="true" pageId="14" pageNumber="99">Lecythis poiteaui</emphasis>
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clade
</emphasis>
(72% BS; 
<figureCitation id="1322E9CDFFDCFFCEFD9F69B0514D8581" box="[526,624,160,185]" captionStart="FIGURE 2" captionStartId="7.[136,229,1995,2016]" captionTargetBox="[152,1424,537,1947]" captionTargetId="figure-166@7.[151,1436,482,1980]" captionTargetPageId="7" captionText="FIGURE 2A. Strict consensus of 66 most parsimonious (MP) trees based on total evidence. Bootstrap values (&gt;50%) are given above the branches.All clades in this figure are part of the Bertholletia clade. The Lecythis pisonis, L. ollaria, Bertholletia excelsa, L. poiteaui, and Corythophora clades of the Bertholletia clade are shown." figureDoi="http://doi.org/10.5281/zenodo.13638675" httpUri="https://zenodo.org/record/13638675/files/figure.png" pageId="14" pageNumber="99">Figs. 2A</figureCitation>
, 
<figureCitation id="1322E9CDFFDCFFCEFDED69B051B68581" box="[636,651,160,185]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">6</figureCitation>
)
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<paragraph id="8BA6F548FFDCFFCEFF1969F85258877D" blockId="14.[136,1452,232,941]" pageId="14" pageNumber="99">
This clade is found from central to eastern Amazonia and disjunct in the coastal forests of eastern 
<collectingCountry id="F30EB5D8FFDCFFCEFA9369F8567B8439" box="[1282,1350,232,257]" name="Brazil" pageId="14" pageNumber="99">Brazil</collectingCountry>
(
<bibRefCitation id="EF8888B9FFDCFFCEFAC969F853F8841D" author="Huang, Y. - Y." pageId="14" pageNumber="99" refId="ref17990" refString="Huang, Y. - Y. (2010) Systematics of Lecythidoideae (Lecythidaceae): with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis. A dissertation submitted to the Graduate Faculty in Biology in partial fullfilment of the requirements for the degree of Doctor of Philosophy, The City University of New York, 140 pp." type="book" year="2010">Huang, 2010</bibRefCitation>
). It is sister to the 
<taxonomicName id="4C198ECBFFDCFFCEFE05681C52C3841D" authorityName="Linnaeus [1759:" authorityYear="1071" box="[404,510,268,293]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="ollaria">
<emphasis id="B96D295AFFDCFFCEFE05681C52C3841D" box="[404,510,268,293]" italics="true" pageId="14" pageNumber="99">L. ollaria</emphasis>
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clade but the five species recovered in it lack the single coil (character 26, 
<figureCitation id="1322E9CDFFDCFFCEFAC1681C53E48471" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6C, D, F, K</figureCitation>
) of that clade. None of the coded morphological characters provide synapomorphies for the 
<taxonomicName id="4C198ECBFFDCFFCEFB786820565E8471" authorityName="O. Berg" authorityYear="1858" box="[1257,1379,304,329]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="poiteaui">
<emphasis id="B96D295AFFDCFFCEFB786820565E8471" box="[1257,1379,304,329]" italics="true" pageId="14" pageNumber="99">L. poiteaui</emphasis>
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clade, but its members possess a long, oblique or geniculate (
<figureCitation id="1322E9CDFFDCFFCEFD7F6844507A8455" box="[750,839,340,365]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6G</figureCitation>
) instead of an erect style; roundish (
<figureCitation id="1322E9CDFFDCFFCEFB71684456708455" box="[1248,1357,340,365]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6J, P</figureCitation>
) instead of longer than broad seeds of the 
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<emphasis id="B96D295AFFDCFFCEFE69686851B884A9" box="[504,645,376,401]" italics="true" pageId="14" pageNumber="99">L. chartacea</emphasis>
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clade (e.g., 
<figureCitation id="1322E9CDFFDCFFCEFC9A686850BC84A9" box="[779,897,376,401]" captionStart="FIGURE 9" captionStartId="19.[136,229,1737,1758]" captionTargetBox="[257,1340,185,1714]" captionTargetId="figure-14@19.[225,1362,175,1726]" captionTargetPageId="19" captionText="FIGURE 9. Seeds of the Lecythis ollaria (A–C), L. chartacea (D–L), and Lecythis poiteaui (M–P) clades. Lecythis ollaria clade—A. L. ollaria (Davidse &amp; González 12096A). B. L. minor (Prance 23172). C. L. tuyrana (not vouchered). All species in this clade have a tendency toward more-or-less fusiform seeds, plane primary veins, major veins oriented along length of seeds, and higher order veins absent, i.e., the areas between the major veins are smooth. Most of the veins of L. ollaria and L. minor only extend for part of the length of the seeds while those of L. tuyrana extend from the base to the apex of the seed. Lecythis chartacea clade—D. L. chartacea (Nee &amp; Mori 4199). E. L. alutacea (Redden et al. 1732). F. L. brancoensis (Silva 48). G. Lecythis rorida (Mori et al. 20428). H. L. retusa (Ramos s.n. NY barcode 00684323). I. L. gracieana (Freitas et al. 745). J. Lecythis parvifructa (Freitas et al. 726). K. L. holcogyne (Mori &amp; Pipoly 15493). L. Eschweilera. simiorum (Clark 4333). Most species in this clade have fusiform or, less frequently, globose seeds, salient primary veins, major veins oriented along the entire length of the seeds, and salient higher order veins. The seeds associated with indehiscent fruits (e.g., those of L. gracieana (I) and L. parvifructa (J), tend to be more globose. Lecythis poiteaui clade—M. L. barnebyi (Costich &amp; dos Santos 836). N. Lecythis ibiriba (Carvalho et al. 6026). O. L. lurida (Prance 26574). P. L. prancei (Mori 20286). All species in this clade" figureDoi="http://doi.org/10.5281/zenodo.13638699" httpUri="https://zenodo.org/record/13638699/files/figure.png" pageId="14" pageNumber="99">Fig. 9D, E</figureCitation>
); dendritic seed venation (
<figureCitation id="1322E9CDFFDCFFCEFB376868561E84A9" box="[1190,1315,376,401]" captionStart="FIGURE 9" captionStartId="19.[136,229,1737,1758]" captionTargetBox="[257,1340,185,1714]" captionTargetId="figure-14@19.[225,1362,175,1726]" captionTargetPageId="19" captionText="FIGURE 9. Seeds of the Lecythis ollaria (A–C), L. chartacea (D–L), and Lecythis poiteaui (M–P) clades. Lecythis ollaria clade—A. L. ollaria (Davidse &amp; González 12096A). B. L. minor (Prance 23172). C. L. tuyrana (not vouchered). All species in this clade have a tendency toward more-or-less fusiform seeds, plane primary veins, major veins oriented along length of seeds, and higher order veins absent, i.e., the areas between the major veins are smooth. Most of the veins of L. ollaria and L. minor only extend for part of the length of the seeds while those of L. tuyrana extend from the base to the apex of the seed. Lecythis chartacea clade—D. L. chartacea (Nee &amp; Mori 4199). E. L. alutacea (Redden et al. 1732). F. L. brancoensis (Silva 48). G. Lecythis rorida (Mori et al. 20428). H. L. retusa (Ramos s.n. NY barcode 00684323). I. L. gracieana (Freitas et al. 745). J. Lecythis parvifructa (Freitas et al. 726). K. L. holcogyne (Mori &amp; Pipoly 15493). L. Eschweilera. simiorum (Clark 4333). Most species in this clade have fusiform or, less frequently, globose seeds, salient primary veins, major veins oriented along the entire length of the seeds, and salient higher order veins. The seeds associated with indehiscent fruits (e.g., those of L. gracieana (I) and L. parvifructa (J), tend to be more globose. Lecythis poiteaui clade—M. L. barnebyi (Costich &amp; dos Santos 836). N. Lecythis ibiriba (Carvalho et al. 6026). O. L. lurida (Prance 26574). P. L. prancei (Mori 20286). All species in this clade" figureDoi="http://doi.org/10.5281/zenodo.13638699" httpUri="https://zenodo.org/record/13638699/files/figure.png" pageId="14" pageNumber="99">Fig. 9M–P</figureCitation>
); and absent (
<figureCitation id="1322E9CDFFDCFFCEFF01688C53E2848D" box="[144,223,412,437]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6J</figureCitation>
) or vestigial aril (
<figureCitation id="1322E9CDFFDCFFCEFE39688C52C1848D" box="[424,508,412,437]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6P</figureCitation>
) versus a more developed basal aril of other clades (characters 47 and 48; 
<figureCitation id="1322E9CDFFDCFFCEFAD5688C569A848D" box="[1348,1447,412,437]" captionStart="FIGURE 4" captionStartId="12.[136,229,1744,1765]" captionTargetBox="[252,1337,266,1683]" captionTargetId="figure-32@12.[210,1378,266,1726]" captionTargetPageId="12" captionText="FIGURE 4. The Lecythis pisonis clade (see Fig. 101 in Mori &amp; Prance, 1990 for vouchers). A. Flower with an open androecial hood. B. Medial longitudinal section of flower with flat androecial hood. The androecial hood possesses a proximal group of staminodes for about one-quarter of its length (not distinguishable in this image) and vestigial stamens for the rest of the length. C. Medial longitudinal section of ovary. Note the stylar collar located just under the stigma. D. Cross-section of 4-locular ovary. E. Operculum with a woody columella that projects into the fruit. F. Fruit base. The fruits of this clade are the largest of all Lecythidaceae. G. Sulcate seed with a long basal aril. Note the funicle projecting from the bottom of the aril. Drawing by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638687" httpUri="https://zenodo.org/record/13638687/files/figure.png" pageId="14" pageNumber="99">Figs. 4G</figureCitation>
, 
<figureCitation id="1322E9CDFFDCFFCEFF1968D0539B84E1" box="[136,166,448,473]" captionStart="FIGURE 5" captionStartId="13.[136,229,1724,1745]" captionTargetBox="[269,1300,307,1679]" captionTargetId="figure-43@13.[266,1322,266,1706]" captionTargetPageId="13" captionText="FIGURE 5. The Lecythis ollaria clade. A–F. L. minor (A based on Galdames 5768; B–F. based on Nee &amp; Mori 3580). G–J. L. tuyrana (based on Galdames 5766). A. Medial longitudinal section of flower showing the single coil and the vestigial stamens limited to the exterior of the coil. B. Apical view of dehisced fruit. C. Lateral view of fruit. D. Basal view of operculum. E. Apical view of operculum. F. Seeds with basal arils. G. Medial longitudinal section of flower showing single but short coil. H. Apical view of fruit with operculum removed and showing seeds inside. I. Basal view of operculum. Note that the columella is not developed. J. Seed with basal aril (above). Note funicle protruding from aril. Photos A and G–J by C. Galdames and photos B–F by S. A. Mori." figureDoi="http://doi.org/10.5281/zenodo.13638689" httpUri="https://zenodo.org/record/13638689/files/figure.png" pageId="14" pageNumber="99">5F</figureCitation>
, 
<figureCitation id="1322E9CDFFDCFFCEFF2268D053C484E1" box="[179,249,448,473]" captionStart="FIGURE 9" captionStartId="19.[136,229,1737,1758]" captionTargetBox="[257,1340,185,1714]" captionTargetId="figure-14@19.[225,1362,175,1726]" captionTargetPageId="19" captionText="FIGURE 9. Seeds of the Lecythis ollaria (A–C), L. chartacea (D–L), and Lecythis poiteaui (M–P) clades. Lecythis ollaria clade—A. L. ollaria (Davidse &amp; González 12096A). B. L. minor (Prance 23172). C. L. tuyrana (not vouchered). All species in this clade have a tendency toward more-or-less fusiform seeds, plane primary veins, major veins oriented along length of seeds, and higher order veins absent, i.e., the areas between the major veins are smooth. Most of the veins of L. ollaria and L. minor only extend for part of the length of the seeds while those of L. tuyrana extend from the base to the apex of the seed. Lecythis chartacea clade—D. L. chartacea (Nee &amp; Mori 4199). E. L. alutacea (Redden et al. 1732). F. L. brancoensis (Silva 48). G. Lecythis rorida (Mori et al. 20428). H. L. retusa (Ramos s.n. NY barcode 00684323). I. L. gracieana (Freitas et al. 745). J. Lecythis parvifructa (Freitas et al. 726). K. L. holcogyne (Mori &amp; Pipoly 15493). L. Eschweilera. simiorum (Clark 4333). Most species in this clade have fusiform or, less frequently, globose seeds, salient primary veins, major veins oriented along the entire length of the seeds, and salient higher order veins. The seeds associated with indehiscent fruits (e.g., those of L. gracieana (I) and L. parvifructa (J), tend to be more globose. Lecythis poiteaui clade—M. L. barnebyi (Costich &amp; dos Santos 836). N. Lecythis ibiriba (Carvalho et al. 6026). O. L. lurida (Prance 26574). P. L. prancei (Mori 20286). All species in this clade" figureDoi="http://doi.org/10.5281/zenodo.13638699" httpUri="https://zenodo.org/record/13638699/files/figure.png" pageId="14" pageNumber="99">9A, B</figureCitation>
). The species of this clade generally have the androecial hood closed (= closed androecium, character 33, 
<figureCitation id="1322E9CDFFDCFFCEFF1968F4521B84C5" box="[136,294,484,509]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Figs. 6D, F, K</figureCitation>
) and petals that are tightly pressed against the androecium, presumably to stop entry into the flowers by non-pollinators. In addition the entrance into the apex of androecial hood is yellow, a color that usually directs bees to a pollinator reward.
</paragraph>
</subSubSection>
<subSubSection id="C303A6C3FFDCFFCEFF2C6B40571A8695" pageId="14" pageNumber="99" type="description">
<paragraph id="8BA6F548FFDCFFCEFF2C6B40571A8695" blockId="14.[136,1452,232,941]" pageId="14" pageNumber="99">
Species of the 
<taxonomicName id="4C198ECBFFDCFFCEFEF56B40511B8751" authorityName="O. Berg" authorityYear="1858" box="[356,550,592,617]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="poiteaui">
<emphasis id="B96D295AFFDCFFCEFEF56B40511B8751" box="[356,550,592,617]" italics="true" pageId="14" pageNumber="99">Lecythis poiteaui</emphasis>
</taxonomicName>
clade that are not bee pollinated are 
<taxonomicName id="4C198ECBFFDCFFCEFC586B4056918751" authority="S. A. Mori (1981 a: 360)" authorityName="S. A. Mori" authorityPageNumber="360" authorityYear="1981" box="[969,1452,592,617]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="barnebyi">
<emphasis id="B96D295AFFDCFFCEFC586B4057AE8751" box="[969,1171,592,617]" italics="true" pageId="14" pageNumber="99">Lecythis barnebyi</emphasis>
S. A. 
<bibRefCitation id="EF8888B9FFDCFFCEFB4B6B4056918751" author="Mori, S. A." box="[1242,1452,592,617]" pageId="14" pageNumber="99" pagination="357 - 370" refId="ref18578" refString="Mori, S. A. (1981 a) New species and combinations in neotropical Lecythidaceae. Brittonia 33: 357 - 370. http: // dx. doi. org / 10.2307 / 2806426" type="journal article" year="1981">Mori (1981a: 360)</bibRefCitation>
</taxonomicName>
(
<figureCitation id="1322E9CDFFDCFFCEFF016B6453D187B5" box="[144,236,628,653]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6A</figureCitation>
) and 
<taxonomicName id="4C198ECBFFDCFFCEFEA36B64529287B5" authorityName="O. Berg" authorityYear="1858" box="[306,431,628,653]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="poiteaui">
<emphasis id="B96D295AFFDCFFCEFEA36B64529287B5" box="[306,431,628,653]" italics="true" pageId="14" pageNumber="99">L. poiteaui</emphasis>
</taxonomicName>
. These two species are nocturnal and bats have been observed taking nectar from their flowers; thus, they are presumed to be bat-pollinated (
<bibRefCitation id="EF8888B9FFDCFFCEFD7D6B8850DE8789" author="Prance, G. T. &amp; Mori, S. A." box="[748,995,664,689]" pageId="14" pageNumber="99" pagination="177 - 181" refId="ref20266" refString="Prance, G. T. &amp; Mori, S. A. (1990) Eschweilera section Jugastrum. In: Mori, S. A. &amp; Prance, G. T. (Eds.) Lecythidaceae - Part II. The zygomorphic-flowered New World genera (Couroupita, Corythophora, Bertholletia, Couratari, Eschweilera, &amp; Lecythis). Flora Neotropica Monograph 21: 177 - 181." type="journal article" year="1990">Mori &amp; Prance, 1990</bibRefCitation>
). These two species also possess similar cuticular papillae on the abaxial leaf blade surface (character 5), a massive number of stamens (character 34; 
<figureCitation id="1322E9CDFFDCFFCEFADE6BAC53A087C1" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6A, B</figureCitation>
), open androecia (character 33; 
<figureCitation id="1322E9CDFFDCFFCEFD976BF051B787C1" box="[518,650,736,761]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Figs. 6A, B</figureCitation>
), petals not pressed against the androecium (
<figureCitation id="1322E9CDFFDCFFCEFB136BF057E187C1" box="[1154,1244,736,761]" captionStart="FIGURE 6" captionStartId="15.[136,229,1732,1753]" captionTargetBox="[241,1358,197,1686]" captionTargetId="figure-13@15.[216,1372,175,1722]" captionTargetPageId="15" captionText="FIGURE 6. The Lecythis poiteaui clade. A–C. L. barnebyi (A–B see Mori &amp; Lepsch-Cunha, 1995 for vouchers and C see Mori 1990c for voucher). D–E. L. prancei (see Mori, 1990c for vouchers). F–J. L. lurida (see Mori, 1990c for vouchers). K–L. L. ibiriba (K–L based on Popovkin 496, M based on Popovkin 497, N–P based on Cardoso 2338). A. Anterior view of flower showing open androecial hood. B. Lateral view of flower showing open androecial hood. C. Medial section of androecium. Note that innermost hood appendages arise from slight expansion of hood, there is a long anterior hood extension, and all of the appendages are swept inward. D. Medial section of androecium. Note that the appendages are swept inward. E. Cross-section of ovary showing mucilage ducts in the ovary wall. F. Medial section of flower. Note that hood appendages are swept inward. G. Medial section of calyx and ovary. Note that style is obliquely oriented and long. H. Cross-section of ovary. Note mucilage ducts in the calyx-lobes. I. Indehiscent fruit, operculum facing downward. J. Seed with dendritic venation and without well-developed aril. K. Medial section of flower. Note that hood appendages are swept inward. L. Cross-section of ovary showing mucilage ducts in ovary wall. M. Lateral view of fruit. N.Apical view of open fruit with seeds inside. O. Basal view of operculum. P. Seed with dendritic venation and poorly-developed aril. Drawings A–B by A. Tangerini and all others by B. Angell." figureDoi="http://doi.org/10.5281/zenodo.13638691" httpUri="https://zenodo.org/record/13638691/files/figure.png" pageId="14" pageNumber="99">Fig. 6A</figureCitation>
), and the presence of at least some anthers (or possibly antherodes) on the hood (character 32). 
<bibRefCitation id="EF8888B9FFDCFFCEFC7B6A1457BE8625" author="Mori, S. A." box="[1002,1155,772,797]" pageId="14" pageNumber="99" pagination="267 - 326" refId="ref18822" refString="Mori, S. A. (1990 b) Lecythis. In: Mori, S. A. &amp; Prance, G. T. (Eds.) Lecythidaceae - Part II. The zygomorphic-flowered New World genera (Couroupita, Corythophora, Couratari, Eschweilera, Lecythis). Flora Neotropica Monograph 21: 267 - 326." type="journal article" year="1990">Mori (1990b)</bibRefCitation>
placed 
<taxonomicName id="4C198ECBFFDCFFCEFB4A6A1451608679" authority="(R. Knuth 1939: 84) S. A. Mori (1981 a: 359)" authorityName="S. A. Mori" authorityPageNumber="359" authorityYear="1981" baseAuthorityName="R. Knuth" baseAuthorityPageNumber="84" baseAuthorityYear="1939" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="brancoensis">
<emphasis id="B96D295AFFDCFFCEFB4A6A1456BD8625" box="[1243,1408,772,797]" italics="true" pageId="14" pageNumber="99">L. brancoensis</emphasis>
(R. 
<bibRefCitation id="EF8888B9FFDCFFCEFF196A3852078679" author="Knuth, R. G. P." box="[136,314,808,833]" pageId="14" pageNumber="99" pagination="1 - 146" refId="ref18185" refString="Knuth, R. G. P. (1939) Lecythidaceae. In: Engler, A. (Ed.) Das Pflanzenreich, Heft 105 (IV, 219 a): 1 - 146." type="book chapter" year="1939">Knuth 1939: 84</bibRefCitation>
) S. A. 
<bibRefCitation id="EF8888B9FFDCFFCEFE1B6A3851608679" author="Mori, S. A." box="[394,605,808,833]" pageId="14" pageNumber="99" pagination="357 - 370" refId="ref18578" refString="Mori, S. A. (1981 a) New species and combinations in neotropical Lecythidaceae. Brittonia 33: 357 - 370. http: // dx. doi. org / 10.2307 / 2806426" type="journal article" year="1981">Mori (1981a: 359)</bibRefCitation>
</taxonomicName>
, along with the two other bat-pollinated species, in 
<taxonomicName id="4C198ECBFFDCFFCEFB376A3856998679" authorityName="O. Berg" authorityYear="1858" box="[1190,1444,808,833]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="section" section="Poiteaui">
<emphasis id="B96D295AFFDCFFCEFB376A38563E8679" box="[1190,1283,808,833]" italics="true" pageId="14" pageNumber="99">Lecythis</emphasis>
sect. 
<emphasis id="B96D295AFFDCFFCEFAD46A3856998679" box="[1349,1444,808,833]" italics="true" pageId="14" pageNumber="99">Poiteaui</emphasis>
</taxonomicName>
, and this relationship was supported by 
<bibRefCitation id="EF8888B9FFDCFFCEFDA86A5C5029865D" author="Huang, Y. &amp; Mori, S. A. &amp; Kelly, L. M." box="[569,788,844,869]" pageId="14" pageNumber="99" pagination="396 - 417" refId="ref18052" refString="Huang, Y. - Y, Mori, S. A. &amp; Kelly, L. M. (2011) A morphological cladistic analysis of Lecythidoideae with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis. Brittonia 63 (3): 396 - 417. http: // dx. doi. org / 10.1007 / s 12228 - 011 - 9202 - 4" type="journal article" year="2011">
Huang 
<emphasis id="B96D295AFFDCFFCEFD1B6A5C5187865D" box="[650,698,844,869]" italics="true" pageId="14" pageNumber="99">et al</emphasis>
. (2011)
</bibRefCitation>
. In contrast, this study places 
<taxonomicName id="4C198ECBFFDCFFCEFBCE6A5C563E865D" authorityName="S. A. Mori" authorityYear="1981" baseAuthorityName="R. Knuth" baseAuthorityYear="1939" box="[1119,1283,844,869]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="brancoensis">
<emphasis id="B96D295AFFDCFFCEFBCE6A5C563E865D" box="[1119,1283,844,869]" italics="true" pageId="14" pageNumber="99">L. brancoensis</emphasis>
</taxonomicName>
in the 
<taxonomicName id="4C198ECBFFDCFFCEFADF6A5C53C586B1" authorityName="O. Berg" authorityYear="1856" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="chartacea">
<emphasis id="B96D295AFFDCFFCEFADF6A5C53C586B1" italics="true" pageId="14" pageNumber="99">Lecythis chartacea</emphasis>
</taxonomicName>
clade. Thus, if 
<taxonomicName id="4C198ECBFFDCFFCEFE276A60516286B1" authorityName="S. A. Mori" authorityYear="1981" baseAuthorityName="R. Knuth" baseAuthorityYear="1939" box="[438,607,880,905]" class="Magnoliopsida" family="Lecythidaceae" genus="Lecythis" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="species" species="brancoensis">
<emphasis id="B96D295AFFDCFFCEFE276A60516286B1" box="[438,607,880,905]" italics="true" pageId="14" pageNumber="99">L. brancoensis</emphasis>
</taxonomicName>
is found to be bat-pollinated as suggested by 
<bibRefCitation id="EF8888B9FFDCFFCEFB156A60561F86B1" author="Mori, S. A." box="[1156,1314,880,905]" pageId="14" pageNumber="99" pagination="267 - 326" refId="ref18822" refString="Mori, S. A. (1990 b) Lecythis. In: Mori, S. A. &amp; Prance, G. T. (Eds.) Lecythidaceae - Part II. The zygomorphic-flowered New World genera (Couroupita, Corythophora, Couratari, Eschweilera, Lecythis). Flora Neotropica Monograph 21: 267 - 326." type="journal article" year="1990">Mori (1990b)</bibRefCitation>
, our results indicate that bat pollination may have evolved twice in New World 
<taxonomicName id="4C198ECBFFDCFFCEFCEC6A84571E8695" box="[893,1059,916,941]" class="Magnoliopsida" family="Lecythidaceae" kingdom="Plantae" order="Ericales" pageId="14" pageNumber="99" phylum="Tracheophyta" rank="family">Lecythidaceae</taxonomicName>
.
</paragraph>
</subSubSection>
</treatment>
</document>