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<mods:title id="30CE7E00BA409A6F0433DC76CC648F84">A molecular phylogeny of Calenduleae (Asteraceae) supports the transfer of Dimorphotheca polyptera DC. to Osteospermum L.</mods:title>
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<treatment id="B91BC43F5713E363FF92117FFDB431FC" LSID="urn:lsid:plazi:treatment:B91BC43F5713E363FF92117FFDB431FC" httpUri="http://treatment.plazi.org/id/B91BC43F5713E363FF92117FFDB431FC" lastPageId="10" lastPageNumber="243" pageId="9" pageNumber="243">
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4.2.
<taxonomicName id="F6B20EAA5713E360FF67117FFEB23203" ID-CoL="45N7" authority="Vaill. ex Moench" box="[145,289,798,818]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">Dimorphotheca</taxonomicName>
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The current circumscription of the genus
<taxonomicName id="F6B20EAA5713E360FDC31137FD56325B" box="[565,709,854,874]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FDC31137FD56325B" box="[565,709,854,874]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
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comprises three main taxonomic entities that were segregated by
<bibRefCitation id="552308D85713E360FD231115FF713293" author="Norlindh, T." pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh, 1943</bibRefCitation>
). The first of these is the original, narrower circumscription of the genus,
<emphasis id="03C6A93B5713E360FE7011CBFD1C328F" box="[390,655,938,958]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FE7011CBFDDC328F" box="[390,591,938,958]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sensu">
Dimorphotheca
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stricto
</emphasis>
, defined by the production of dimorphic ray and disc fruits (e.g.,
<figureCitation id="A98969AC5713E360FD8011A6FD5F32EB" box="[630,716,967,986]" captionStart="Fig" captionStartId="1.[100,125,1798,1812]" captionTargetBox="[234,1354,148,1767]" captionTargetId="figure-1@1.[234,1354,148,1768]" captionTargetPageId="1" captionText="Fig. 1. Morphologicial diversity in Calenduleae. a,b — Garuleum pinnatifidum pinnatisect leaves and blue-violet rays; c — Osteospermum moniliferum unripe drupaceous fruits developing from the ray florets of the central capitulum; d —fruit from a single individual within Osteospermum sect. Calendula; e — Garuleum bipinnatum, fruiting head showing dimorphic fruit developing from ray and disc florets; f — Garuleum pinnatifidum, fruiting head showing fruit developing only from rays; g — Dimorphotheca cuneata capitulum; h —fruiting head of a species from Osteospermum sect. Calendula; i —Dimorphotheca polyptera fruiting head showing winged ray fruit and flattened disc fruit; j — Osteospermum nordenstamii capitulum; k — D. sinuata immature ray and disk fruits showing packing; l — D. pluvialis ray (left) and disc (right) fruit; m — O. sinuatum, unripe three-winged fruit with apical air cavity; n — D. cuneata capitula; o — O.connatum capitula; p — O. scariosum capitulum; q — D. sinuata capitulum from below. Photo credits: a,b — Gerhard Malan, iNaturalist; c — Joan Young, NaturesWow; d — Fig. 27B from Gon¸calves et al. (2018) showing Calendula suffruticosa subsp. suffruticosa; e — Thabo Maphisa, OperationWildflower; f — R. Sadler; g — Liz Makings; h —Plantas y Hongos website: Calendula arvensis; i — N.van Berkel, iNaturalist; j,k,m,n,o,q — N.Bergh; l — K.Pillay; p — A.G.Ellis." figureDoi="http://doi.org/10.5281/zenodo.10496952" httpUri="https://zenodo.org/record/10496952/files/figure.png" pageId="9" pageNumber="243">Fig. 1k, l</figureCitation>
; taxa indicated by “(D)” after the taxon names in
<figureCitation id="A98969AC5713E360FDE81182FDC732C7" box="[542,596,995,1014]" captionStart="Fig" captionStartId="7.[100,125,1716,1730]" captionTargetBox="[256,1331,148,1685]" captionTargetId="figure-97@7.[256,1331,148,1686]" captionTargetPageId="7" captionText="Fig. 4. Bayesian consensus tree, as in Fig.3, but with all nodes collapsed except those with BS Ż 75% and/or PP Ż 0.9. Type species are marked with an asterisk in front of the species name. Characters were obtained from taxonomic literature cited in the text.Floral, life-history, fruit and leaf characters for Osteospermum sect.Calendula were obtained from Meikle (1976). Disc style characters were obtained from Norlindh (1943). Seed oil characters were obtained from Smith et al. (1960), Earle et al. (1962), Barclay and Earle (1965), and (Chisholm and Hopkins, 1967). Letters in parentheses after species names indicate taxonomic placement: C — Osteospermum sect. Calendula; Ca — former Castalis; D — Dimorphotheca sensu stricto; H — O. sect. Homocarpae; M — former Monoculus; O — O. sect. Coriaceae; OsB — O. sect.Blaxium; P — O. sect. Polygalinae; R — O. sect. Trialatae; T — O. subgenus. Tripteris (former Tripteris)." figureDoi="http://doi.org/10.5281/zenodo.10496962" httpUri="https://zenodo.org/record/10496962/files/figure.png" pageId="9" pageNumber="243">Fig. 4</figureCitation>
), and comprising only seven species, three of which are represented in our tree (
<figureCitation id="A98969AC5713E360FD31119EFD693523" box="[711,762,1023,1042]" captionStart="Fig" captionStartId="7.[100,125,1716,1730]" captionTargetBox="[256,1331,148,1685]" captionTargetId="figure-97@7.[256,1331,148,1686]" captionTargetPageId="7" captionText="Fig. 4. Bayesian consensus tree, as in Fig.3, but with all nodes collapsed except those with BS Ż 75% and/or PP Ż 0.9. Type species are marked with an asterisk in front of the species name. Characters were obtained from taxonomic literature cited in the text.Floral, life-history, fruit and leaf characters for Osteospermum sect.Calendula were obtained from Meikle (1976). Disc style characters were obtained from Norlindh (1943). Seed oil characters were obtained from Smith et al. (1960), Earle et al. (1962), Barclay and Earle (1965), and (Chisholm and Hopkins, 1967). Letters in parentheses after species names indicate taxonomic placement: C — Osteospermum sect. Calendula; Ca — former Castalis; D — Dimorphotheca sensu stricto; H — O. sect. Homocarpae; M — former Monoculus; O — O. sect. Coriaceae; OsB — O. sect.Blaxium; P — O. sect. Polygalinae; R — O. sect. Trialatae; T — O. subgenus. Tripteris (former Tripteris)." figureDoi="http://doi.org/10.5281/zenodo.10496962" httpUri="https://zenodo.org/record/10496962/files/figure.png" pageId="9" pageNumber="243">Fig. 4</figureCitation>
;
<emphasis id="03C6A93B5713E360FF921678FEAA351C" box="[100,313,1049,1069]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FF921678FF5D351C" box="[100,206,1049,1069]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="pluvialis">D. pluvialis</taxonomicName>
,
<taxonomicName id="F6B20EAA5713E360FF2F1678FEAA351C" box="[217,313,1049,1069]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sinuata">D. sinuata</taxonomicName>
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, and
<taxonomicName id="F6B20EAA5713E360FE861678FE77351C" box="[368,484,1049,1069]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FE861678FE77351C" box="[368,484,1049,1069]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
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). The second entity is
<taxonomicName id="F6B20EAA5713E360FD331678FEF33578" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="section" section="Blaxium">
<emphasis id="03C6A93B5713E360FD331678FF283578" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
section
<emphasis id="03C6A93B5713E360FEE21654FEF33578" box="[276,352,1077,1097]" italics="true" pageId="9" pageNumber="243">Blaxium</emphasis>
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, placed by
<bibRefCitation id="552308D85713E360FE2E1657FDEB3578" author="Norlindh, T." box="[472,632,1078,1097]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
in
<taxonomicName id="F6B20EAA5713E360FD681654FF073554" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FD681654FF073554" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
since all species produce only ray fruit, and comprising eight species. Three of these are included in our tree, viz.
<taxonomicName id="F6B20EAA5713E360FDAF160CFF1B35AC" authority="E. Phillips" authorityName="E. Phillips" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="jucunda">
<emphasis id="03C6A93B5713E360FDAF160CFD2E35B0" box="[601,701,1133,1153]" italics="true" pageId="9" pageNumber="243">D. jucunda</emphasis>
E.Phillips
</taxonomicName>
,
<taxonomicName id="F6B20EAA5713E360FF6416E8FEB335AC" authority="DC." authorityName="DC." box="[146,288,1161,1181]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="ecklonis">
<emphasis id="03C6A93B5713E360FF6416E8FF6535AC" box="[146,246,1161,1181]" italics="true" pageId="9" pageNumber="243">D. ecklonis</emphasis>
DC.
</taxonomicName>
, and
<taxonomicName id="F6B20EAA5713E360FEA216E8FE2C35AC" authorityName="Breitwieser" authorityYear="2191" box="[340,447,1161,1181]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="fruticosa">
<emphasis id="03C6A93B5713E360FEA216E8FE2C35AC" box="[340,447,1161,1181]" italics="true" pageId="9" pageNumber="243">D. fruticosa</emphasis>
</taxonomicName>
(L.) DC, and indicated by “(OsB)” after the taxon names in
<figureCitation id="A98969AC5713E360FEAF16C7FE1F3588" box="[345,396,1190,1209]" captionStart="Fig" captionStartId="7.[100,125,1716,1730]" captionTargetBox="[256,1331,148,1685]" captionTargetId="figure-97@7.[256,1331,148,1686]" captionTargetPageId="7" captionText="Fig. 4. Bayesian consensus tree, as in Fig.3, but with all nodes collapsed except those with BS Ż 75% and/or PP Ż 0.9. Type species are marked with an asterisk in front of the species name. Characters were obtained from taxonomic literature cited in the text.Floral, life-history, fruit and leaf characters for Osteospermum sect.Calendula were obtained from Meikle (1976). Disc style characters were obtained from Norlindh (1943). Seed oil characters were obtained from Smith et al. (1960), Earle et al. (1962), Barclay and Earle (1965), and (Chisholm and Hopkins, 1967). Letters in parentheses after species names indicate taxonomic placement: C — Osteospermum sect. Calendula; Ca — former Castalis; D — Dimorphotheca sensu stricto; H — O. sect. Homocarpae; M — former Monoculus; O — O. sect. Coriaceae; OsB — O. sect.Blaxium; P — O. sect. Polygalinae; R — O. sect. Trialatae; T — O. subgenus. Tripteris (former Tripteris)." figureDoi="http://doi.org/10.5281/zenodo.10496962" httpUri="https://zenodo.org/record/10496962/files/figure.png" pageId="9" pageNumber="243">Fig. 4</figureCitation>
. The third taxonomic entity is the former genus
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<emphasis id="03C6A93B5713E360FF2D16A0FEB135E4" box="[219,290,1217,1237]" italics="true" pageId="9" pageNumber="243">Castalis</emphasis>
</taxonomicName>
, comprising just three species and segregated from
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<emphasis id="03C6A93B5713E360FF6A16BCFEBF35C0" box="[156,300,1245,1265]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
by
<bibRefCitation id="552308D85713E360FEA016BFFE6635C0" author="Norlindh, T." box="[342,501,1246,1265]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
on the basis of sterile ray florets, a unique feature in
<taxonomicName id="F6B20EAA5713E360FE9C169BFE71343C" box="[362,482,1274,1293]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="245" phylum="Tracheophyta" rank="tribe" tribe="Calenduleae">Calenduleae</taxonomicName>
.
<taxonomicName id="F6B20EAA5713E360FE1D1698FDA1343C" box="[491,562,1273,1293]" class="Magnoliopsida" family="Asteraceae" genus="Castalis" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FE1D1698FDA1343C" box="[491,562,1273,1293]" italics="true" pageId="9" pageNumber="243">Castalis</emphasis>
</taxonomicName>
is represented in our tree by the single species
<taxonomicName id="F6B20EAA5713E360FE841775FE5B3419" box="[370,456,1300,1320]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="tragus">
<emphasis id="03C6A93B5713E360FE841775FE5B3419" box="[370,456,1300,1320]" italics="true" pageId="9" pageNumber="243">C. tragus</emphasis>
</taxonomicName>
(indicated by “(Ca)” in
<figureCitation id="A98969AC5713E360FD481774FD663418" box="[702,757,1301,1321]" captionStart="Fig" captionStartId="7.[100,125,1716,1730]" captionTargetBox="[256,1331,148,1685]" captionTargetId="figure-97@7.[256,1331,148,1686]" captionTargetPageId="7" captionText="Fig. 4. Bayesian consensus tree, as in Fig.3, but with all nodes collapsed except those with BS Ż 75% and/or PP Ż 0.9. Type species are marked with an asterisk in front of the species name. Characters were obtained from taxonomic literature cited in the text.Floral, life-history, fruit and leaf characters for Osteospermum sect.Calendula were obtained from Meikle (1976). Disc style characters were obtained from Norlindh (1943). Seed oil characters were obtained from Smith et al. (1960), Earle et al. (1962), Barclay and Earle (1965), and (Chisholm and Hopkins, 1967). Letters in parentheses after species names indicate taxonomic placement: C — Osteospermum sect. Calendula; Ca — former Castalis; D — Dimorphotheca sensu stricto; H — O. sect. Homocarpae; M — former Monoculus; O — O. sect. Coriaceae; OsB — O. sect.Blaxium; P — O. sect. Polygalinae; R — O. sect. Trialatae; T — O. subgenus. Tripteris (former Tripteris)." figureDoi="http://doi.org/10.5281/zenodo.10496962" httpUri="https://zenodo.org/record/10496962/files/figure.png" pageId="9" pageNumber="243">Fig. 4</figureCitation>
). Lastly,
<taxonomicName id="F6B20EAA5713E360FF511751FE093474" authority="(Thunb.) Harv." authorityName="Harv." baseAuthorityName="Thunb." box="[167,410,1328,1349]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="pinnata">
<emphasis id="03C6A93B5713E360FF511751FE943475" box="[167,263,1328,1348]" italics="true" pageId="9" pageNumber="243">D. pinnata</emphasis>
(Thunb.) Harv.
</taxonomicName>
was transferred to
<taxonomicName id="F6B20EAA5713E360FDA31751FD763475" box="[597,741,1328,1348]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FDA31751FD763475" box="[597,741,1328,1348]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
by Nordenstam (2006). This is a morphologically unusual annual species with pinnatisect leaves and spiny ray fruits, placed by
<bibRefCitation id="552308D85713E360FD5D1708FF3734A9" author="Norlindh, T." pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
in the monospecific section
<emphasis id="03C6A93B5713E360FE4017E5FE5734A9" box="[438,452,1412,1432]" italics="true" pageId="9" pageNumber="243">O</emphasis>
. sect.
<taxonomicName id="F6B20EAA5713E360FE0917E5FD6A34A9" authority="(DC.) T. Norl" authorityName="T. Norl" baseAuthorityName="DC." box="[511,761,1412,1432]" class="Magnoliopsida" family="Asteraceae" genus="Acanthotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FE0917E5FDED34A9" box="[511,638,1412,1432]" italics="true" pageId="9" pageNumber="243">Acanthotheca</emphasis>
(DC.) T. Norl
</taxonomicName>
.. However, we were unable to include a specimen of
<taxonomicName id="F6B20EAA5713E360FDA917C1FD523485" box="[607,705,1440,1460]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="pinnata">
<emphasis id="03C6A93B5713E360FDA917C1FD523485" box="[607,705,1440,1460]" italics="true" pageId="9" pageNumber="243">D. pinnata</emphasis>
</taxonomicName>
in our tree, and it has never been examined for seed oil composition. Despite this lack of evidence regarding its generic position,
<taxonomicName id="F6B20EAA5713E360FD5717B9FC9134DD" box="[673,770,1496,1516]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="pinnata">
<emphasis id="03C6A93B5713E360FD5717B9FC9134DD" box="[673,770,1496,1516]" italics="true" pageId="9" pageNumber="243">D. pinnata</emphasis>
</taxonomicName>
most closely resembles the annual
<emphasis id="03C6A93B5713E360FE351795FD433739" box="[451,720,1524,1544]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FE351795FD1D3739" box="[451,654,1524,1544]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sensu">
Dimorphotheca
<taxonomicNameLabel id="18F514405713E360FDAC1795FD1D3739" box="[602,654,1524,1544]" pageId="9" pageNumber="243" sensu="stricto">sensu</taxonomicNameLabel>
</taxonomicName>
stricto
</emphasis>
species in floral characters.
</paragraph>
<paragraph id="310D75295713E360FF72144DFA163331" blockId="9.[100,771,854,1990]" lastBlockId="9.[818,1488,157,1990]" pageId="9" pageNumber="243">
Several authors acknowledged strong similarities of involucre and floral characters, disc fruit structure, and phytochemistry, between
<emphasis id="03C6A93B5713E360FF921402FE173746" box="[100,388,1635,1655]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FF921402FEAA3746" box="[100,313,1635,1655]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sensu">
Dimorphotheca
<taxonomicNameLabel id="18F514405713E360FEF31402FEAA3746" box="[261,313,1635,1655]" pageId="9" pageNumber="243" sensu="stricto">sensu</taxonomicNameLabel>
</taxonomicName>
stricto
</emphasis>
and the other taxonomic entities described above (
<bibRefCitation id="552308D85713E360FEED14E1FE2237A2" author="Candolle, A. P." box="[283,433,1664,1683]" pageId="9" pageNumber="243" refId="ref10862" refString="Candolle, A. P., 1838. Prodromus Systematis Naturalis Regni Vegetabilis VI. Treuttel et W urtz €, Paris." type="book" year="1838">Candolle, 1838</bibRefCitation>
;
<bibRefCitation id="552308D85713E360FE3514E0FDD937A2" author="Harvey, W. H." box="[451,586,1664,1684]" pageId="9" pageNumber="243" pagination="44 - 530" refId="ref11347" refString="Harvey, W. H., 1865. Compositae. in: Harvey, W. H., Sonder, O. W. (Eds) Flora Capensis. Hodges, Smith &amp; Co., Dublin, pp. 44 - 530. vol. 3." type="journal article" year="1865">Harvey, 1865</bibRefCitation>
;
<bibRefCitation id="552308D85713E360FDAB14E1FD6637A2" author="Norlindh, T." box="[605,757,1664,1683]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh, 1943</bibRefCitation>
). However,
<bibRefCitation id="552308D85713E360FF3114FDFEC9379E" author="Norlindh, T." box="[199,346,1692,1711]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943</bibRefCitation>
,
<bibRefCitation id="552308D85713E360FE9314FDFE05379E" author="Norlindh, T." box="[357,406,1692,1711]" pageId="9" pageNumber="243" pagination="961 - 987" refId="ref12042" refString="Norlindh, T., 1978. Ch. 34: calenduleae - systematic review. In: Heywood, V. H., Harborne, J. B., Turner, B. L. (Eds.), Biology and Chemistry of the Compositae. Academic Press, Cornell, pp. 961 - 987." type="book chapter" year="1978">1978</bibRefCitation>
) was reluctant to expand the concept of
<taxonomicName id="F6B20EAA5713E360FF8914D6FE9C37FA" box="[127,271,1719,1739]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FF8914D6FE9C37FA" box="[127,271,1719,1739]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
to incorporate them as this would have removed any clear morphological synapomorphies for the genus.
<bibRefCitation id="552308D85713E360FD5D14B5FF373632" author="Norlindh, T." pageId="9" pageNumber="243" pagination="961 - 987" refId="ref12042" refString="Norlindh, T., 1978. Ch. 34: calenduleae - systematic review. In: Heywood, V. H., Harborne, J. B., Turner, B. L. (Eds.), Biology and Chemistry of the Compositae. Academic Press, Cornell, pp. 961 - 987." type="book chapter" year="1978">Norlindh (1978)</bibRefCitation>
instead discussed resurrecting the genus
<taxonomicName id="F6B20EAA5713E360FDC1148EFD293632" authority="Cass." authorityName="Cass." box="[567,698,1775,1795]" class="Magnoliopsida" family="Asteraceae" genus="Blaxium" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FDC1148EFD103632" box="[567,643,1775,1795]" italics="true" pageId="9" pageNumber="243">Blaxium</emphasis>
Cass.
</taxonomicName>
, clearly intending to maintain three separate genera for the species currently in
<taxonomicName id="F6B20EAA5713E360FF761547FE83360B" box="[128,272,1830,1850]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FF761547FE83360B" box="[128,272,1830,1850]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
. However,
<bibRefCitation id="552308D85713E360FE721546FDC1360A" author="Nordenstam, B." box="[388,594,1831,1851]" pageId="9" pageNumber="243" pagination="46 - 49" refId="ref11668" refString="Nordenstam, B., 1994 b. New combinations in the Calenduleae. Compos. Newsl. 25, 46 - 49." type="journal article" year="1994">Nordenstam (1994b)</bibRefCitation>
considered these entities to be congeneric on the basis of a morphological, cytological and chemical parsimony analysis, and transferred both
<taxonomicName id="F6B20EAA5713E360FD68153FFE8A36BF" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="section" section="Blaxium">
<emphasis id="03C6A93B5713E360FD68153FFF0736BF" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
sect.
<emphasis id="03C6A93B5713E360FF3B151BFE8A36BF" box="[205,281,1914,1934]" italics="true" pageId="9" pageNumber="243">Blaxium</emphasis>
</taxonomicName>
and
<taxonomicName id="F6B20EAA5713E360FEBC151BFE0236BF" box="[330,401,1914,1934]" class="Magnoliopsida" family="Asteraceae" genus="Castalis" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FEBC151BFE0236BF" box="[330,401,1914,1934]" italics="true" pageId="9" pageNumber="243">Castalis</emphasis>
</taxonomicName>
to
<taxonomicName id="F6B20EAA5713E360FE44151BFDD136BF" box="[434,578,1914,1934]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FE44151BFDD136BF" box="[434,578,1914,1934]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
, thereby nearly tripling the number of species. Together with the reversal by Nordenstam (2006) of the earlier transfer of
<taxonomicName id="F6B20EAA5713E360FD9815D3FD4F36F7" box="[622,732,1970,1990]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="pinnata">
<emphasis id="03C6A93B5713E360FD9815D3FD4F36F7" box="[622,732,1970,1990]" italics="true" pageId="9" pageNumber="243">D. pinnata</emphasis>
</taxonomicName>
to
<taxonomicName id="F6B20EAA5713E360FCC412FCFC2E3180" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[818,957,157,177]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FCC412FCFC2E3180" box="[818,957,157,177]" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
by
<bibRefCitation id="552308D85713E360FC1812FFFB023180" author="Norlindh, T." box="[1006,1169,158,177]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
, this constitutes the currently accepted circumscription of
<taxonomicName id="F6B20EAA5713E360FBBE12D8FCF931D8" authority="(Manning and Goldblatt, 2012)" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FBBE12D8FB4B31FC" box="[1096,1240,185,205]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
(
<bibRefCitation id="552308D85713E360FB1312DBFCF731D8" author="Manning, J. C. &amp; Goldblatt, P." pageId="9" pageNumber="243" refId="ref11473" refString="Manning, J. C., Goldblatt, P., 2012. Plants of the greater cape floristic region 1: the core cape flora. Strelitzia 29, SANBI, Pretoria." type="book" year="2012">Manning and Goldblatt, 2012</bibRefCitation>
)
</taxonomicName>
. With the sole exception of
<taxonomicName id="F6B20EAA5713E360FB7412B4FB6631D8" box="[1154,1269,213,233]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FB7412B4FB6631D8" box="[1154,1269,213,233]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
, the enlarged circumscription of
<taxonomicName id="F6B20EAA5713E360FC431291FBD63035" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[949,1093,240,260]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FC431291FBD63035" box="[949,1093,240,260]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
proposed by
<bibRefCitation id="552308D85713E360FB161293FA213034" author="Nordenstam, B." box="[1248,1458,242,261]" pageId="9" pageNumber="243" pagination="46 - 49" refId="ref11668" refString="Nordenstam, B., 1994 b. New combinations in the Calenduleae. Compos. Newsl. 25, 46 - 49." type="journal article" year="1994">Nordenstam (1994b)</bibRefCitation>
is strongly supported in our tree, despite the lack of clear morphological characters defining the genus. Moreover, although only seven of the » 20 currently accepted species of
<taxonomicName id="F6B20EAA5713E360FB751325FA803069" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[1155,1299,324,344]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FB751325FA803069" box="[1155,1299,324,344]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
are included in our analysis, these represent all three of the main historical taxonomic entities now included in the genus. The strongest un-contradicted synapomorphy for the genus is the presence of dimorphecolic acid in the seed oils, since this has been found in only twelve of many investigated species of
<taxonomicName id="F6B20EAA5713E360FC2B13B0FBC730D5" box="[989,1108,465,484]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="245" phylum="Tracheophyta" rank="tribe" tribe="Calenduleae">Calenduleae</taxonomicName>
, all of which are
<taxonomicName id="F6B20EAA5713E360FB0313B1FA5C30D5" box="[1269,1487,464,484]" pageId="9" pageNumber="243">
<emphasis id="03C6A93B5713E360FB0313B1FA1630D5" box="[1269,1413,464,484]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
species
</taxonomicName>
(
<bibRefCitation id="552308D85713E360FCCF138DFC7230CE" author="Smith, Jr., C. R. &amp; Wilson, T. L. &amp; Melvin, E. H. &amp; Wolff, I. A." box="[825,993,492,512]" pageId="9" pageNumber="243" pagination="1417 - 1421" refId="ref12413" refString="Smith, Jr., C. R., Wilson, T. L., Melvin, E. H., Wolff, I. A., 1960. Dimorphecolic acid - a unique hydroxydienoid fatty acid. J. Am. Chem. Soc. 83 (6), 1417 - 1421." type="journal article" year="1960">Smith et al., 1960</bibRefCitation>
;
<bibRefCitation id="552308D85713E360FC1B138DFB1830CE" author="Earle, F. R. &amp; Wolff, I. A. &amp; Glass, C. A. &amp; Jones, Q." box="[1005,1163,492,512]" pageId="9" pageNumber="243" pagination="381 - 383" refId="ref10961" refString="Earle, F. R., Wolff, I. A., Glass, C. A., Jones, Q., 1962. Search for new industrial oils. VII. J. Americal Oil Chemist. Soc. 39, 381 - 383." type="journal article" year="1962">Earle et al., 1962</bibRefCitation>
;
<bibRefCitation id="552308D85713E360FB61138DFAEB30CE" author="Barclay, A. S. &amp; Earle, F. R." box="[1175,1400,492,511]" pageId="9" pageNumber="243" pagination="33 - 43" refId="ref10628" refString="Barclay, A. S., Earle, F. R., 1965. The search for new industrial crops V. The South African Calenduleae (Compositae) as a source of new oil seeds. Econ. Bot. 19, 33 - 43." type="journal article" year="1965">Barclay and Earle, 1965</bibRefCitation>
).
</paragraph>
<paragraph id="310D75295713E360FCA71066FBB737A2" blockId="9.[818,1488,157,1990]" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FCA71066FBDC332A" ID-CoL="6CVS8" box="[849,1103,519,539]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FCA71066FBDC332A" box="[849,1103,519,539]" italics="true" pageId="9" pageNumber="243">Dimorphotheca polyptera</emphasis>
</taxonomicName>
was included without question in
<emphasis id="03C6A93B5713E360FCC41042FBD83306" box="[818,1099,547,567]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FCC41042FB973306" box="[818,1028,547,567]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sensu">
Dimorphotheca
<taxonomicNameLabel id="18F514405713E360FC391042FB973306" box="[975,1028,547,567]" pageId="9" pageNumber="243" sensu="stricto">sensu</taxonomicNameLabel>
</taxonomicName>
stricto
</emphasis>
by both
<bibRefCitation id="552308D85713E360FB461045FAC63306" author="Norlindh, T." box="[1200,1365,548,567]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
and earlier workers (e.g.
<bibRefCitation id="552308D85713E360FC321021FBCE3362" author="Candolle, A. P." box="[964,1117,576,595]" pageId="9" pageNumber="243" refId="ref10862" refString="Candolle, A. P., 1838. Prodromus Systematis Naturalis Regni Vegetabilis VI. Treuttel et W urtz €, Paris." type="book" year="1838">Candolle, 1838</bibRefCitation>
;
<bibRefCitation id="552308D85713E360FB861020FB6B3362" author="Harvey, W. H." box="[1136,1272,576,596]" pageId="9" pageNumber="243" pagination="44 - 530" refId="ref11347" refString="Harvey, W. H., 1865. Compositae. in: Harvey, W. H., Sonder, O. W. (Eds) Flora Capensis. Hodges, Smith &amp; Co., Dublin, pp. 44 - 530. vol. 3." type="journal article" year="1865">Harvey, 1865</bibRefCitation>
) because it has the defining characteristic of the genus, producing dimorphic fruits from both disc and ray florets.
<bibRefCitation id="552308D85713E360FB751019FABB33BA" author="Norlindh, T." box="[1155,1320,632,651]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
noted however that
<taxonomicName id="F6B20EAA5713E360FC9510F2FC4F3396" ID-CoL="6CVS8" box="[867,988,659,679]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FC9510F2FC4F3396" box="[867,988,659,679]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
differs from all other
<taxonomicName id="F6B20EAA5713E360FB3F10F2FA3E3396" box="[1225,1453,659,679]" pageId="9" pageNumber="243">
<emphasis id="03C6A93B5713E360FB3F10F2FACF3396" box="[1225,1372,659,679]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
species
</taxonomicName>
by its short, yellow ray ligules. In this respect, it is more similar to many members of
<taxonomicName id="F6B20EAA5713E360FC0110AAFA9A33EE" box="[1015,1289,715,735]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="subSpecies" species="sensu" subSpecies="amplo">
<emphasis id="03C6A93B5713E360FC0110AAFA9A33EE" box="[1015,1289,715,735]" italics="true" pageId="9" pageNumber="243">Osteospermum sensu amplo</emphasis>
</taxonomicName>
. The structure and size of the disc fruits produced by
<taxonomicName id="F6B20EAA5713E360FB401086FAA233CA" ID-CoL="6CVS8" box="[1206,1329,743,763]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FB401086FAA233CA" box="[1206,1329,743,763]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
are similar to those of both
<taxonomicName id="F6B20EAA5713E360FC221163FBF73227" ID-CoL="3FBV9" box="[980,1124,770,790]" class="Magnoliopsida" family="Asteraceae" genus="Garuleum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="bipinnatum">
<emphasis id="03C6A93B5713E360FC221163FBF73227" box="[980,1124,770,790]" italics="true" pageId="9" pageNumber="243">G. bipinnatum</emphasis>
</taxonomicName>
and
<taxonomicName id="F6B20EAA5713E360FB501163FA8E3227" ID-CoL="6CVRV" box="[1190,1309,770,790]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="montana">
<emphasis id="03C6A93B5713E360FB501163FA8E3227" box="[1190,1309,770,790]" italics="true" pageId="9" pageNumber="243">D. montana</emphasis>
</taxonomicName>
, being obovoidoblong rather than distinctly cordate, and their small size (
<quantity id="F64AD8CC5713E360FA511141FCFE327E" metricMagnitude="-3" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="9" pageNumber="243" unit="mm" value="3.5" valueMax="4.0" valueMin="3.0">
3
<emphasis id="03C6A93B5713E360FA49117EFA5C3203" box="[1471,1487,799,818]" italics="true" pageId="9" pageNumber="243"></emphasis>
4 mm
</quantity>
long) is similar to those of both
<taxonomicName id="F6B20EAA5713E360FB36115BFAB1327F" ID-CoL="6CW48" box="[1216,1314,826,846]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="sinuata">
<emphasis id="03C6A93B5713E360FB36115BFAB1327F" box="[1216,1314,826,846]" italics="true" pageId="9" pageNumber="243">D. sinuata</emphasis>
</taxonomicName>
(
<quantity id="F64AD8CC5713E360FAC4115AFA08327F" box="[1330,1435,827,846]" metricMagnitude="-3" metricUnit="m" metricValue="3.5" metricValueMax="4.0" metricValueMin="3.0" pageId="9" pageNumber="243" unit="mm" value="3.5" valueMax="4.0" valueMin="3.0">
3
<emphasis id="03C6A93B5713E360FAB0115AFAC5327F" box="[1350,1366,827,846]" italics="true" pageId="9" pageNumber="243"></emphasis>
4 mm
</quantity>
) and
<taxonomicName id="F6B20EAA5713E360FCC41137FC2C325B" ID-CoL="3FBV9" box="[818,959,854,874]" class="Magnoliopsida" family="Asteraceae" genus="Garuleum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="bipinnatum">
<emphasis id="03C6A93B5713E360FCC41137FC2C325B" box="[818,959,854,874]" italics="true" pageId="9" pageNumber="243">G. bipinnatum</emphasis>
</taxonomicName>
(
<quantity id="F64AD8CC5713E360FC241136FBDB325B" box="[978,1096,855,874]" metricMagnitude="-3" metricUnit="m" metricValue="4.5" metricValueMax="5.0" metricValueMin="4.0" pageId="9" pageNumber="243" unit="mm" value="4.5" valueMax="5.0" valueMin="4.0">
4
<emphasis id="03C6A93B5713E360FC1D1136FC68325B" box="[1003,1019,855,874]" italics="true" pageId="9" pageNumber="243"></emphasis>
5 mm
</quantity>
), but smaller than most
<taxonomicName id="F6B20EAA5713E360FAA01136FA5C325B" box="[1366,1487,855,874]" class="Magnoliopsida" family="Asteraceae" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="245" phylum="Tracheophyta" rank="tribe" tribe="Calenduleae">Calenduleae</taxonomicName>
disc fruits, which can be up to
<quantity id="F64AD8CC5713E360FB871112FB2A32B7" box="[1137,1209,883,902]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="9" pageNumber="243" unit="mm" value="20.0">20 mm</quantity>
long (e.g. in
<taxonomicName id="F6B20EAA5713E360FAB51113FA5132B7" ID-CoL="6D7RW" box="[1347,1474,882,902]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="spectabilis">
<emphasis id="03C6A93B5713E360FAB51113FA5132B7" box="[1347,1474,882,902]" italics="true" pageId="9" pageNumber="243">D. spectabilis</emphasis>
</taxonomicName>
). The ray fruits in
<taxonomicName id="F6B20EAA5713E360FC0F11EFFBE53293" ID-CoL="6CVS8" box="[1017,1142,910,930]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FC0F11EFFBE53293" box="[1017,1142,910,930]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
are polymorphic, and Candolle (1837) distinguished three different shapes of the ray fruits, establishing a novel section
<taxonomicName id="F6B20EAA5713E360FBA311A7FB5F32EB" box="[1109,1228,966,986]" pageId="9" pageNumber="243" rank="section" section="Triplocarpae">
<emphasis id="03C6A93B5713E360FBA311A7FB5F32EB" box="[1109,1228,966,986]" italics="true" pageId="9" pageNumber="243">Triplocarpae</emphasis>
</taxonomicName>
within
<taxonomicName id="F6B20EAA5713E360FAE911A7FA2132EB" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[1311,1458,966,986]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FAE911A7FA2132EB" box="[1311,1458,966,986]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
to accommodate it. The winged ray fruits of
<taxonomicName id="F6B20EAA5713E360FB011183FAE232C7" ID-CoL="6CVS8" box="[1271,1393,994,1014]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FB011183FAE232C7" box="[1271,1393,994,1014]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
are pigmented on the body, terete to (more commonly) trigonous, and with varying degrees of development of pale wing-like processes on the angles (
<figureCitation id="A98969AC5713E360FC221657FB853578" box="[980,1046,1078,1097]" captionStart="Fig" captionStartId="1.[100,125,1798,1812]" captionTargetBox="[234,1354,148,1767]" captionTargetId="figure-1@1.[234,1354,148,1768]" captionTargetPageId="1" captionText="Fig. 1. Morphologicial diversity in Calenduleae. a,b — Garuleum pinnatifidum pinnatisect leaves and blue-violet rays; c — Osteospermum moniliferum unripe drupaceous fruits developing from the ray florets of the central capitulum; d —fruit from a single individual within Osteospermum sect. Calendula; e — Garuleum bipinnatum, fruiting head showing dimorphic fruit developing from ray and disc florets; f — Garuleum pinnatifidum, fruiting head showing fruit developing only from rays; g — Dimorphotheca cuneata capitulum; h —fruiting head of a species from Osteospermum sect. Calendula; i —Dimorphotheca polyptera fruiting head showing winged ray fruit and flattened disc fruit; j — Osteospermum nordenstamii capitulum; k — D. sinuata immature ray and disk fruits showing packing; l — D. pluvialis ray (left) and disc (right) fruit; m — O. sinuatum, unripe three-winged fruit with apical air cavity; n — D. cuneata capitula; o — O.connatum capitula; p — O. scariosum capitulum; q — D. sinuata capitulum from below. Photo credits: a,b — Gerhard Malan, iNaturalist; c — Joan Young, NaturesWow; d — Fig. 27B from Gon¸calves et al. (2018) showing Calendula suffruticosa subsp. suffruticosa; e — Thabo Maphisa, OperationWildflower; f — R. Sadler; g — Liz Makings; h —Plantas y Hongos website: Calendula arvensis; i — N.van Berkel, iNaturalist; j,k,m,n,o,q — N.Bergh; l — K.Pillay; p — A.G.Ellis." figureDoi="http://doi.org/10.5281/zenodo.10496952" httpUri="https://zenodo.org/record/10496952/files/figure.png" pageId="9" pageNumber="243">Fig. 1i</figureCitation>
). Three-winged ray fruits are common in both
<taxonomicName id="F6B20EAA5713E360FC991630FB913554" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[879,1026,1105,1125]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FC991630FB913554" box="[879,1026,1105,1125]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
and
<taxonomicName id="F6B20EAA5713E360FBB31630FB403554" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[1093,1235,1105,1125]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FBB31630FB403554" box="[1093,1235,1105,1125]" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
(
<figureCitation id="A98969AC5713E360FB1C1633FABB3554" box="[1258,1320,1106,1125]" captionStart="Fig" captionStartId="7.[100,125,1716,1730]" captionTargetBox="[256,1331,148,1685]" captionTargetId="figure-97@7.[256,1331,148,1686]" captionTargetPageId="7" captionText="Fig. 4. Bayesian consensus tree, as in Fig.3, but with all nodes collapsed except those with BS Ż 75% and/or PP Ż 0.9. Type species are marked with an asterisk in front of the species name. Characters were obtained from taxonomic literature cited in the text.Floral, life-history, fruit and leaf characters for Osteospermum sect.Calendula were obtained from Meikle (1976). Disc style characters were obtained from Norlindh (1943). Seed oil characters were obtained from Smith et al. (1960), Earle et al. (1962), Barclay and Earle (1965), and (Chisholm and Hopkins, 1967). Letters in parentheses after species names indicate taxonomic placement: C — Osteospermum sect. Calendula; Ca — former Castalis; D — Dimorphotheca sensu stricto; H — O. sect. Homocarpae; M — former Monoculus; O — O. sect. Coriaceae; OsB — O. sect.Blaxium; P — O. sect. Polygalinae; R — O. sect. Trialatae; T — O. subgenus. Tripteris (former Tripteris)." figureDoi="http://doi.org/10.5281/zenodo.10496962" httpUri="https://zenodo.org/record/10496962/files/figure.png" pageId="9" pageNumber="243">Fig. 4</figureCitation>
), but they are rarely discolorous, and do not possess broad, pale wings divided into processes. This
<typeStatus id="EE09CB8B5713E360FC0816EAFBB935AF" box="[1022,1066,1163,1182]" pageId="9" pageNumber="243">type</typeStatus>
of fruit is, however, found in one species of
<taxonomicName id="F6B20EAA5713E360FCBA16C4FC493588" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[844,986,1189,1209]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FCBA16C4FC493588" box="[844,986,1189,1209]" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
, viz.
<taxonomicName id="F6B20EAA5713E360FBE616C4FA823588" ID-CoL="6TBFJ" authority="(Compton) Norl" authorityName="Norl" baseAuthorityName="Compton" box="[1040,1297,1189,1209]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="bolusii">
<emphasis id="03C6A93B5713E360FBE616C4FBF93588" box="[1040,1130,1189,1209]" italics="true" pageId="9" pageNumber="243">O. bolusii</emphasis>
(Compton) Norl
</taxonomicName>
, which was placed by
<bibRefCitation id="552308D85713E360FCA216A3FC6B35E4" author="Norlindh, T." box="[852,1016,1218,1237]" pageId="9" pageNumber="243" refId="ref11896" refString="Norlindh, T., 1943. Studies in the Calenduleae I. Monograph of the Genera Dimorphotheca, Castalis, Osteospermum, Gibbaria and Chrysanthemoides. C. W. K. Gleerup, Lund." type="book" year="1943">Norlindh (1943)</bibRefCitation>
and
<bibRefCitation id="552308D85713E360FBC716A3FAF935E4" author="Manning, J. C. &amp; Goldblatt, P." box="[1073,1386,1218,1237]" pageId="9" pageNumber="243" refId="ref11473" refString="Manning, J. C., Goldblatt, P., 2012. Plants of the greater cape floristic region 1: the core cape flora. Strelitzia 29, SANBI, Pretoria." type="book" year="2012">Manning and Goldblatt (2012)</bibRefCitation>
in
<taxonomicName id="F6B20EAA5713E360FA6716A0FB0B35C0" authority="Norl. While" authorityName="Norl. While" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="section" section="Trialatae">
<emphasis id="03C6A93B5713E360FA6716A0FC1835C0" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
sect.
<emphasis id="03C6A93B5713E360FC3116BCFB8835C0" box="[967,1051,1245,1265]" italics="true" pageId="9" pageNumber="243">Trialatae</emphasis>
Norl. While
</taxonomicName>
<taxonomicName id="F6B20EAA5713E360FB5616BCFB6935C0" ID-CoL="6TBFJ" box="[1184,1274,1245,1265]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="bolusii">
<emphasis id="03C6A93B5713E360FB5616BCFB6935C0" box="[1184,1274,1245,1265]" italics="true" pageId="9" pageNumber="243">O. bolusii</emphasis>
</taxonomicName>
differs from
<taxonomicName id="F6B20EAA5713E360FA7416BCFCF7343C" ID-CoL="6CVS8" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FA7416BCFCF7343C" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
in being a perennial shrublet with entire leaf margins, in having female-sterile disc florets, and in having monomorphic ray fruits, the two species have strong floral similarities, their ray ligules being short, narrow, and uniformly pale yellow. In features of the fruits, ray florets, and in phylogenetic placement,
<taxonomicName id="F6B20EAA5713E360FA761709FCF734A9" ID-CoL="6CVS8" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FA761709FCF734A9" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
does not conform to
<taxonomicName id="F6B20EAA5713E360FBB417E5FB4634A9" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[1090,1237,1412,1432]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FBB417E5FB4634A9" box="[1090,1237,1412,1432]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
and agrees most closely with species in
<taxonomicName id="F6B20EAA5713E360FC1617C1FB6B3485" box="[992,1272,1440,1460]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="subSpecies" species="sensu" subSpecies="amplo">
<emphasis id="03C6A93B5713E360FC1617C1FB6B3485" box="[992,1272,1440,1460]" italics="true" pageId="9" pageNumber="243">Osteospermum sensu amplo</emphasis>
</taxonomicName>
, and particularly in sect.
<emphasis id="03C6A93B5713E360FC9E17DDFC2F34E1" box="[872,956,1468,1488]" italics="true" pageId="9" pageNumber="243">Trialatae</emphasis>
. As far as we are aware, the phytochemistry of
<taxonomicName id="F6B20EAA5713E360FA4A17DDFC1E34DD" ID-CoL="6CVS8" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FA4A17DDFC1E34DD" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
has never been investigated but we predict that it is unlikely to contain dimorphecolic acid. While elucidation of the closest relatives of
<taxonomicName id="F6B20EAA5713E360FBF11471FB103715" ID-CoL="6CVS8" box="[1031,1155,1552,1572]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FBF11471FB103715" box="[1031,1155,1552,1572]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
within
<emphasis id="03C6A93B5713E360FB171471FA043715" box="[1249,1431,1552,1572]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FB171471FAFC3715" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[1249,1391,1552,1572]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">Osteospermum</taxonomicName>
s.a
</emphasis>
. will require greater sampling, our analyses confirm that the species is phylogenetically distant from the remainder of
<taxonomicName id="F6B20EAA5713E360FACA1426FA5C376A" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[1340,1487,1607,1627]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FACA1426FA5C376A" box="[1340,1487,1607,1627]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
using multiple accessions from across the geographic range of the species (
<figureCitation id="A98969AC5713E360FC7E14E1FC2D37A2" box="[904,958,1664,1683]" captionStart="Fig" captionStartId="6.[100,125,1537,1551]" captionTargetBox="[258,1338,153,1501]" captionTargetId="figure-245@6.[250,1304,148,1507]" captionTargetPageId="6" captionText="Fig. 3. Bayesian consensus tree with average branch lengths inferred from concatenated nuclear and plastid DNA with associated indels. Numbers at nodes represent parsimony bootstrap percentages followed by Bayesian posterior probability (PP) values (only indicated on nodes receiving Ż 50% bootstrap and Ż 0.50 PP support). Tip name abbreviations: C = Osteospermum sect.Calendula, D = Dimorphotheca, G = Garuleum, O = Osteospermum. Members of Norlindhs subgenus Tripteris are indicated by a symbol indicated three-winged fruits.Orange, bold font indicates the accessions of D. polyptera." figureDoi="http://doi.org/10.5281/zenodo.10496960" httpUri="https://zenodo.org/record/10496960/files/figure.png" pageId="9" pageNumber="243">Fig. 3</figureCitation>
;
<tableCitation id="7C3040925713E360FC3814E1FB8537A2" box="[974,1046,1664,1683]" captionStart="Table 2" captionStartId="4.[117,161,1192,1206]" captionTargetPageId="4" captionText="Table 2 Characteristics of the phylogenetic data.PI&quot; indicates characters that are phylogenetically informative." pageId="9" pageNumber="243">Table 2</tableCitation>
).
</paragraph>
<paragraph id="310D75295713E363FCA714FDFDB431FC" blockId="9.[818,1488,157,1990]" lastBlockId="10.[100,770,157,205]" lastPageId="10" lastPageNumber="244" pageId="9" pageNumber="243">
While the exact phylogenetic position of
<taxonomicName id="F6B20EAA5713E360FAFF14FAFA10379E" ID-CoL="6CVS8" box="[1289,1411,1691,1711]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FAFF14FAFA10379E" box="[1289,1411,1691,1711]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
within
<taxonomicName id="F6B20EAA5713E360FCC414D6FC5337FA" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[818,960,1719,1739]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FCC414D6FC5337FA" box="[818,960,1719,1739]" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
is uncertain, its retention in
<taxonomicName id="F6B20EAA5713E360FB0714D6FA1737FA" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[1265,1412,1719,1739]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FB0714D6FA1737FA" box="[1265,1412,1719,1739]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
clearly renders that genus non-monophyletic. Also, removal of
<taxonomicName id="F6B20EAA5713E360FA8914B2FCF73632" ID-CoL="6CVS8" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FA8914B2FCF73632" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
from
<taxonomicName id="F6B20EAA5713E360FC50148EFBAA3632" ID-CoL="45N7" baseAuthorityName="Manning and Goldblatt" baseAuthorityYear="2012" box="[934,1081,1775,1795]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FC50148EFBAA3632" box="[934,1081,1775,1795]" italics="true" pageId="9" pageNumber="243">Dimorphotheca</emphasis>
</taxonomicName>
simplifies the morphological definition of the genus, since it now contains no species with short, narrow and uniformly pale yellow ray ligules. We accordingly transfer
<taxonomicName id="F6B20EAA5713E360FA4A1547FC1E3667" ID-CoL="6CVS8" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FA4A1547FC1E3667" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
to
<taxonomicName id="F6B20EAA5713E360FC471523FBAC3667" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[945,1087,1858,1878]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5713E360FC471523FBAC3667" box="[945,1087,1858,1878]" italics="true" pageId="9" pageNumber="243">Osteospermum</emphasis>
</taxonomicName>
based on its phylogenetic position, ray ligule morphology, and ray fruit structure, and provisionally assign it to
<emphasis id="03C6A93B5713E360FC41151BFC5636BF" box="[951,965,1914,1934]" italics="true" pageId="9" pageNumber="243">O</emphasis>
. sect.
<emphasis id="03C6A93B5713E360FBE6151BFBF736BF" box="[1040,1124,1914,1934]" italics="true" pageId="9" pageNumber="243">Trialatae</emphasis>
. While
<taxonomicName id="F6B20EAA5713E360FB49151BFAA936BF" ID-CoL="6CVS8" box="[1215,1338,1914,1934]" class="Magnoliopsida" family="Asteraceae" genus="Dimorphotheca" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="species" species="polyptera">
<emphasis id="03C6A93B5713E360FB49151BFAA936BF" box="[1215,1338,1914,1934]" italics="true" pageId="9" pageNumber="243">D. polyptera</emphasis>
</taxonomicName>
is anomalous within
<emphasis id="03C6A93B5713E360FC8815F7FBA5369B" box="[894,1078,1942,1962]" italics="true" pageId="9" pageNumber="243">
<taxonomicName id="F6B20EAA5713E360FC8815F7FB9F369B" ID-CoL="63GQQ" authorityName="sensu (Manning and Goldblatt" authorityYear="2012" box="[894,1036,1942,1962]" class="Magnoliopsida" family="Asteraceae" genus="Osteospermum" kingdom="Plantae" order="Asterales" pageId="9" pageNumber="243" phylum="Tracheophyta" rank="genus">Osteospermum</taxonomicName>
s.a.
</emphasis>
in being the only known species with female-fertile disc florets, there is a precedent for such an anomaly in the genus
<taxonomicName id="F6B20EAA5710E363FEBA12FCFE393180" ID-CoL="62SSD" box="[332,426,157,177]" class="Magnoliopsida" family="Asteraceae" genus="Garuleum" kingdom="Plantae" order="Asterales" pageId="10" pageNumber="244" phylum="Tracheophyta" rank="genus">
<emphasis id="03C6A93B5710E363FEBA12FCFE393180" box="[332,426,157,177]" italics="true" pageId="10" pageNumber="244">Garuleum</emphasis>
</taxonomicName>
, where a single species produces disc fruits in addition to the usual ray fruits.
</paragraph>
</subSubSection>
</treatment>
</document>