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<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" article-type="brief-report" xml:lang="en">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Bothalia</journal-id>
<journal-title-group>
<journal-title>Bothalia African Biodiversity &#x0026; Conservation</journal-title>
</journal-title-group>
<issn pub-type="ppub">0006-8241</issn>
<issn pub-type="epub">2311-9284</issn>
<publisher>
<publisher-name>AOSIS</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">ABC-47-2260</article-id>
<article-id pub-id-type="doi">10.4102/abc.v47i1.2260</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Short Communication</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Status of the Genera <italic>Colpoon, Osyris</italic> and <italic>Rhoiacarpos</italic> in South Africa</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<contrib-id contrib-id-type="orcid">http://orcid.org/0000-0001-8519-0517</contrib-id>
<name>
<surname>Nickrent</surname>
<given-names>Daniel L.</given-names>
</name>
<xref ref-type="aff" rid="AF0001">1</xref>
</contrib>
<aff id="AF0001"><label>1</label>College of Science, Department of Plant Biology, Southern Illinois University, United States</aff>
</contrib-group>
<author-notes>
<corresp id="cor1"><bold>Corresponding author:</bold> Daniel Nickrent, <email xlink:href="nickrent@plant.siu.edu">nickrent@plant.siu.edu</email></corresp>
</author-notes>
<pub-date pub-type="epub"><day>13</day><month>11</month><year>2017</year></pub-date>
<pub-date pub-type="collection"><year>2017</year></pub-date>
<volume>47</volume>
<issue>1</issue>
<elocation-id>2260</elocation-id>
<history>
<date date-type="received"><day>16</day><month>05</month><year>2017</year></date>
<date date-type="accepted"><day>10</day><month>10</month><year>2017</year></date>
</history>
<permissions>
<copyright-statement>&#x00A9; 2017. The Authors</copyright-statement>
<copyright-year>2017</copyright-year>
<license license-type="open-access" xlink:href="https://creativecommons.org/licenses/by/4.0/">
<license-p>Licensee: AOSIS. This work is licensed under the Creative Commons Attribution License.</license-p>
</license>
</permissions>
<abstract>
<sec id="st1">
<title>Background</title>
<p>The taxonomic and phylogenetic status of <italic>Colpoon, Osyris</italic> and <italic>Rhoiacarpos</italic> (Santalaceae, Osyrideae) is reviewed.</p>
</sec>
<sec id="st2">
<title>Objectives</title>
<p>To resolve confusion regarding whether <italic>Colpoon</italic> is deserving of generic status separate from <italic>Osyris</italic>.</p>
</sec>
<sec id="st3">
<title>Methods</title>
<p>Existing morphological information was examined for the three genera as well as previously published molecular phylogenies.</p>
</sec>
<sec id="st4">
<title>Results</title>
<p>From both morphological and phylogenetic perspectives, <italic>Colpoon</italic> is distinct from <italic>Osyris</italic>. The status of <italic>Rhoiacarpos</italic> was not contentious and this genus is also easily differentiated from the other two genera in Osyrideae.</p>
</sec>
<sec id="st5">
<title>Conclusions</title>
<p><italic>Colpoon</italic> and <italic>Osyris</italic> are not congeneric; therefore, floras, databases and herbarium collections should recognise these as distinct taxa.</p>
</sec>
</abstract>
</article-meta>
</front>
<body>
<sec id="s0001">
<title>Introduction</title>
<p>Over the past several decades, some confusion has surrounded the taxonomic circumscription of three genera of South African Santalaceae, tribe Osyrideae: <italic>Osyris</italic> L. (1753), <italic>Colpoon</italic> P.J.Bergius (1767) and <italic>Rhoiacarpos</italic> A.DC (1857). The taxonomic history and generic boundaries (from a morphological perspective) of these three taxa were discussed by Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>). For several decades following Stauffer&#x2019;s work, these three genera were generally treated as distinct, as shown in &#x2018;Plants of Southern Africa: Names and Distribution&#x2019; by Arnold and De Wet (<xref ref-type="bibr" rid="CIT0001">1993</xref>). In 1994, Hilliard published a one-page note on <italic>Colpoon</italic> where she followed the treatment in Flora Capensis (Hill <xref ref-type="bibr" rid="CIT0007">1915</xref>) and reduced <italic>Colpoon compressum</italic> P.J.Bergius to a synonym of <italic>Osyris abyssinica</italic> Hochst. ex A.Rich. (= <italic>O. lanceolata</italic> Hochst. &#x0026; Steud.). As stated by Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>), &#x2018;There is, however, no essential differences in floral detail or in the structure of the inflorescence; those given by Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) are illusory&#x2019;. But are the differences between the three genera given in Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>, <xref ref-type="table" rid="T0001">Table 1</xref>) real or imagined? It should be noted that Stauffer examined specimens, including the types, from Zurich, Geneva, Paris, London and Kew. His work was the most comprehensive up to that time and he concluded that the three species &#x2018;can be clearly distinguished morphologically and among themselves show no transitions&#x2019;. Most web sites, recent floras and field guides that appeared after Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>) accepted two genera for South Africa, <italic>Osyris</italic> and <italic>Rhoiacarpos</italic>, with <italic>Colpoon</italic> listed as a synonym of the former. Examples include Goldblatt and Manning (<xref ref-type="bibr" rid="CIT0006">2000</xref>), Bean and Johns (<xref ref-type="bibr" rid="CIT0002">2005</xref>), Germishuizen et al. (<xref ref-type="bibr" rid="CIT0005">2006</xref>), Mucina and Rutherford (<xref ref-type="bibr" rid="CIT0011">2006</xref>) and Manning (<xref ref-type="bibr" rid="CIT0009">2007</xref>). This trend was reversed by Manning and Goldblatt (<xref ref-type="bibr" rid="CIT0010">2012</xref>) who took into account molecular phylogenetic data that showed the three genera were distinct. Despite this publication, the concept of two Osyrideae genera for South Africa persists in herbarium collections and both popular and scientific works. Therefore, the purpose of this paper is to summarise the existing morphological and molecular information that supports recognising three distinct genera.</p>
<table-wrap id="T0001">
<label>TABLE 1</label>
<caption><p>Morphological characters for some Osyrideae.</p></caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th valign="top" align="left">Morphological characters</th>
<th valign="top" align="left"><italic>Osyris lanceolata</italic></th>
<th valign="top" align="left"><italic>Colpoon compressum</italic></th>
<th valign="top" align="left"><italic>Rhoiacarpos capensis</italic></th>
</tr>
</thead>
<tbody>
<tr>
<td align="left">Phylotaxy</td>
<td align="left">alternate (spiral 2/5)</td>
<td align="left">decussate, sometimes subopposite or alternate via displacement</td>
<td align="left">decussate, sometimes subopposite</td>
</tr>
<tr>
<td align="left">Leaf shape</td>
<td align="left">elliptical, acute to cuneate base</td>
<td align="left">elliptical, acute to cuneate base</td>
<td align="left">ovate, cordate base</td>
</tr>
<tr>
<td align="left">Leaf texture</td>
<td align="left">soft</td>
<td align="left">soft</td>
<td align="left">coriaceous</td>
</tr>
<tr>
<td align="left">Leaf surfaces</td>
<td align="left">dull above and below</td>
<td align="left">dull above and below</td>
<td align="left">shiny above, dull below</td>
</tr>
<tr>
<td align="left">Leaf margin</td>
<td align="left">plane</td>
<td align="left">plane</td>
<td align="left">revolute</td>
</tr>
<tr>
<td align="left">Petiole</td>
<td align="left">present</td>
<td align="left">present</td>
<td align="left">very short</td>
</tr>
<tr>
<td align="left">Inflorescence position</td>
<td align="left">axillary</td>
<td align="left">terminal</td>
<td align="left">terminal</td>
</tr>
<tr>
<td align="left">Inflorescence type</td>
<td align="left">unifloral (female), monochasia (male and female), compound and compressed monochasia (male)</td>
<td align="left">paniculate with compressed axes, monochasial</td>
<td align="left">paniculate with compressed axes, monochasial</td>
</tr>
<tr>
<td align="left">Inflorescence bracts</td>
<td align="left">abscising</td>
<td align="left">abscising or persistent below fruit</td>
<td align="left">persistent below fruit</td>
</tr>
<tr>
<td align="left">Plant sex</td>
<td align="left">dioecious (androdioecious?)</td>
<td align="left">flowers bisexual</td>
<td align="left">flowers bisexual</td>
</tr>
<tr>
<td align="left">Petal number</td>
<td align="left">3 (4)</td>
<td align="left">4 (&#x2212;6)</td>
<td align="left">(4) 5 (6)</td>
</tr>
<tr>
<td align="left">Style length</td>
<td align="left">to height of anthers or beyond</td>
<td align="left">nearly absent (stigma sessile)</td>
<td align="left">to height of anthers or beyond</td>
</tr>
<tr>
<td align="left">Disk shape</td>
<td align="left">flat, spherical</td>
<td align="left">flat, lobe number equal to petal number</td>
<td align="left">saucer-shaped, lobe number equal to petal number</td>
</tr>
<tr>
<td align="left">Fruit shape</td>
<td align="left">spheroid to ellipsoid</td>
<td align="left">obovoid</td>
<td align="left">spheroid to ellipsoid</td>
</tr>
<tr>
<td align="left">Petals on fruit apex</td>
<td align="left">not persistent</td>
<td align="left">not persistent</td>
<td align="left">persistent</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn><p><italic>Source</italic>: Author&#x2019;s own work, modified and updated from Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>)</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec id="s0002">
<title>Molecular phylogenetic data</title>
<p>The first comprehensive molecular phylogenetic study of Santalaceae was by Der and Nickrent (<xref ref-type="bibr" rid="CIT0004">2008</xref>) that used nuclear small-subunit ribosomal DNA and chloroplast rbcL and matK. A portion of the tree there referred to as the &#x2018;<italic>Santalum</italic> clade&#x2019; (now Santalaceae s. str.) is shown in <xref ref-type="fig" rid="F0001">Figure 1a</xref>. The clade composed of <italic>Osyris, Nestronia, Rhoiacarpos</italic> and <italic>Colpoon</italic> (tribe Osyrideae) is strongly supported as monophyletic. Moreover, <italic>Osyris quadripartita</italic> (Europe) was not sister to <italic>Colpoon</italic>, with <italic>Rhoiacarpos capensis</italic> occupying that position. It is relevant that Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) also saw this association based on morphology:</p>
<disp-quote>
<p>Therefore, an association of <italic>Colpoon</italic> and <italic>Osyris</italic> can not be accepted. If anything, one finds understanding of the union of <italic>Colpoon</italic> and <italic>Rhoiacarpos</italic> according to the actions of Baillon and Bentham. (p. 392)</p>
</disp-quote>
<fig id="F0001">
<label>FIGURE 1</label>
<caption><p>A. Phylogenetics of Santalaceae <italic>sensu stricto</italic>. Maximum likelihood bootstrap, Bayesian posterior probabilities and maximum parsimony bootstrap values are shown at the nodes where &#x2018;&#x2013;&#x2019; = &#x003C; 50&#x0025;. (a) Portion of the three-gene phylogenetic tree from Der and Nickrent (<xref ref-type="bibr" rid="CIT0004">2008</xref>). (b) Portion of the seven-gene phylogenetic tree from Su et al. (<xref ref-type="bibr" rid="CIT0013">2015</xref>).</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="ABC-47-2260-g001.tif"/>
</fig>
<p>One could argue that because no South African accession of <italic>Osyris</italic> was included, this result should not be used to address generic delimitation in this geographic region. A seven-gene study that sampled nuclear, chloroplast and mitochondrial genes from 197 members of Santalales was published (Su et al. <xref ref-type="bibr" rid="CIT0013">2015</xref>) and a portion of the resulting tree is shown in <xref ref-type="fig" rid="F0001">Figure 1b</xref>. In that study, <italic>Osyris lanceolata</italic> from South Africa was included and it is sister to the European <italic>Osyris quadripartita</italic>. As before, <italic>Osyris</italic> is sister to a clade containing <italic>Nestronia, Rhoiacarpos</italic> and <italic>Colpoon</italic> with the latter two sisters.</p>
</sec>
<sec id="s0003">
<title>Morphological data</title>
<p>A comparison of the vegetative and reproductive morphologies of <italic>Osyris, Colpoon</italic> and <italic>Rhoiacarpos</italic> shows that these genera share a number of features and yet differ in others (<xref ref-type="table" rid="T0001">Table 1</xref> and <xref ref-type="fig" rid="F0002">Figures 2</xref>&#x2013;<xref ref-type="fig" rid="F0004">4</xref>).</p>
<fig id="F0002">
<label>FIGURE 2</label>
<caption><p><italic>Osyris lanceolata</italic> vegetative and reproductive morphology. (a) Habit of plant with male flowers showing abundant axillary inflorescences. (b) Axillary male inflorescences. The left-hand shoot (*) may appear racemose but note that new leafy innovations are present at the apex. Branching of the floral peduncles indicates these units are compound monochasia (cymes). (c) Axillary male inflorescences. Although some appear dichasial (*), the lateral floral buds are of unequal age, thus indicating that these are compound monochasia with compressed axes (cf. Stauffer <xref ref-type="bibr" rid="CIT0012">1961</xref>). (d) Axillary female inflorescences. Note that the flowers occur singly or in two-flowered monochasia. The anthers in these flowers (*) do not appear functional, and hence are interpreted as staminodes. (e) Young fruit in axillary position. Note the recaulescent bracts (*) at a node opposite a leaf. (f) Mature fruit.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="ABC-47-2260-g002.tif"/>
</fig>
<fig id="F0003">
<label>FIGURE 3</label>
<caption><p><italic>Colpoon compressum</italic> vegetative and reproductive morphology. (a) Flowering and fruiting shoots. Note the swollen termini of the panicle peduncles (*) representing compressed axes (cf. Stauffer <xref ref-type="bibr" rid="CIT0012">1961</xref>). (b) Side view of inflorescence showing compressed monochasia, minute deciduous inflorescence bracts and a 3-merous flower (*) among mostly 4-merous ones. (c) Top view of inflorescence showing all 4-merous flowers, floral disks and essentially sessile stigmas. (d) Inflorescence showing persistent bracts along edges of swollen panicle termini and flower abortion. (e) Young vegetative innovation showing metatopic displacement resulting in alternate phylotaxy. (f) Mature fruit occurring at swollen peduncular apex with scars from abortive flowers.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="ABC-47-2260-g003.tif"/>
</fig>
<fig id="F0004">
<label>FIGURE 4</label>
<caption><p><italic>Rhoiacarpos capensis</italic> vegetative and reproductive morphology. (a) Habit of plant in fruit. (b) Shoot with terminal paniculate inflorescences. Note the opposite as well as subopposite (*) leaves. (c) Inflorescence. Note persistent bracts below flowers. (d, e) Closer view of the 5-merous flowers. (f) Young infructescence. Note bracts persisting on the peduncles. (g) Mature fruit showing persistent petals at apex.</p></caption>
<graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="ABC-47-2260-g004.tif"/>
</fig>
<sec id="s20004">
<title>Leaves</title>
<p>All three genera have simple, entire leaves with mucronate apices. All show winged or ridged stems with the ridges merging with leaf petiole bases. Leaf phylotaxy was used by Hill (<xref ref-type="bibr" rid="CIT0007">1915</xref>) to justify merging <italic>Colpoon</italic> with <italic>Osyris</italic> because both alternate and opposite leaves were seen in the former. Variation in phylotaxy was noted by Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) where <italic>C. compressum</italic> was scored as decussate, rarely with the two leaves somewhat shifted. Bean (<xref ref-type="bibr" rid="CIT0003">1990</xref>) scored <italic>C. compressum</italic> as &#x2018;leaves usually opposite to subopposite, less often alternate&#x2019; and the newly described species <italic>Colpoon speciosum</italic> as &#x2018;leaves as often alternate as subopposite, less often opposite&#x2019;. Although <italic>Rhoiacarpos</italic> leaves were scored as opposite by Hill (<xref ref-type="bibr" rid="CIT0007">1915</xref>), it too can show the subopposite condition (<xref ref-type="fig" rid="F0004">Figure 4b</xref>). Thus, phylotaxy appears to be a rather plastic morphological feature in Osyrideae. Shifting (displacement) of leaf position through development can be a form of metatopy (Weberling <xref ref-type="bibr" rid="CIT0015">1989</xref>) possibly influenced by genetic or environmental factors.</p>
<p><italic>Rhoiacarpos</italic> is easily distinguished from the other two genera by leaf features. Its leaves are nearly sessile, bifacial (shiny above and dull below), with cordate bases and somewhat revolute margins. The leaves of <italic>Osyris</italic> and <italic>Colpoon</italic> differ in these characters, but are quite similar to each other overall. Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>) states for both <italic>Osyris</italic> and <italic>Colpoon</italic> &#x2018;both leaf surfaces closely and minutely white-dotted&#x2019;, but this feature has not been observed by this author in the field, herbarium or in photographs.</p>
</sec>
<sec id="s20005">
<title>Inflorescences</title>
<p>The inflorescences of <italic>Osyris</italic> are mainly axillary whereas those of <italic>Colpoon</italic> and <italic>Rhoiacarpos</italic> are terminal. It is interesting that Hill (<xref ref-type="bibr" rid="CIT0007">1915</xref>) describes the inflorescence of <italic>Rhoiacarpos</italic> as a panicle composed of 3-flowered axillary cymules. In <xref ref-type="fig" rid="F0004">Figure 4</xref> of Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>), both <italic>Rhoiacarpos</italic> and <italic>Colpoon</italic> are diagrammed with 3-flowered units. In other Santalales, this 3-flowered cymule unit is equivalent to a dichasium. But the situation in Osyrideae appears more complex, mainly owing to compression and reduction (loss of flowers). Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>) indicates flowers in <italic>Osyris lanceolata</italic> &#x2018;may occur in 2&#x2013;3-flowered dichasia&#x2019;. Technically, a dichasium is 3-flowered whereas a monochasium is 2-flowered. Upon first examination, the inflorescence units may appear to be dichasial such as in <italic>Osyris</italic> (<xref ref-type="fig" rid="F0002">Figure 2c</xref>), <italic>Colpoon</italic> (<xref ref-type="fig" rid="F0003">Figure 3b</xref>), and <italic>Rhoiacarpos</italic> (<xref ref-type="fig" rid="F0004">Figure 4c</xref>), but in these cases the pair of flowers (or buds) subtending the older, terminal flower are unequal in age. Moreover, the older of the two subtending buds is often present as a lateral innovation. For these reasons, it seems best to describe these units as monochasia or compound monochasia. Simple (2-flowered) monochasia are frequently seen in the female inflorescences of <italic>Osyris</italic> (<xref ref-type="fig" rid="F0002">Figure 2d</xref>).</p>
<p>The terminal peduncles in the panicles of <italic>Colpoon</italic> are often swollen (<xref ref-type="fig" rid="F0003">Figure 3a</xref> and <xref ref-type="fig" rid="F0003">d</xref>). In his diagram, Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) coloured black those inflorescence axes that become compressed. Thus, the cluster of flowers seen arising from these swollen apices in <italic>Colpoon</italic> is interpreted as being a series of reduced monochasia (or dichasia according to Stauffer <xref ref-type="bibr" rid="CIT0012">1961</xref>). The phenomenon of syndesmy (incorporation of partial inflorescence units and their axes into the main axis) is well documented in cymoid types resulting in a coenosome (Weberling <xref ref-type="bibr" rid="CIT0015">1989</xref>). Looking at the lower inflorescence branch in <xref ref-type="fig" rid="F0003">Figure 3a</xref>, the six flowers and buds are of different ages: two past anthesis, one at anthesis, one older bud and two younger buds. This situation could be interpreted as two dichasia, but because the ages of the flowers are wrong, this situation might best be interpreted as three monochasia.</p>
<p>The components of the inflorescence axes (peduncles) are often subtended by small bracts. Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) indicated that these abscise in <italic>Osyris</italic>, abscise or are persistent in <italic>Colpoon</italic> and are persistent in <italic>Rhoiacarpos</italic>. The study by Bean (<xref ref-type="bibr" rid="CIT0003">1990</xref>) confirms that the persistence of these bracts in <italic>C. compressum</italic> is polymorphic (compare <xref ref-type="fig" rid="F0003">Figure 3a</xref> and <xref ref-type="fig" rid="F0003">d</xref>) and further shows that in <italic>C. speciosum</italic> the bracts are persistent and enlarge upon fruiting.</p>
</sec>
<sec id="s20006">
<title>Flowers</title>
<p>The flowers in <italic>Colpoon</italic> and <italic>Rhoiacarpos</italic> are bisexual, whereas in <italic>Osyris lanceolata</italic> flowers appear to be unisexual. Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>) stated, &#x2018;the flowers of <italic>Osyris lanceolata</italic> are hermaphrodite and male, the plants being androdioecious&#x2019;, but no evidence was provided that supported the presence of bisexual versus female flowers. Female flowers possess stamens (<xref ref-type="fig" rid="F0002">Figure 2d</xref>), but if the anthers can be shown to not produce pollen, then these are staminodes and the flower is functionally female.</p>
<p>Flower merosity is notoriously variable in Santalaceae, often showing three or four different forms within the same inflorescence. But for differentiating the three genera considered here, this character has some value when the most common number is considered. Thus, <italic>Osyris</italic> generally has 3-merous flowers, occasionally having 4-merous flowers. <italic>Colpoon</italic> typically has 4-merous flowers, but 5- and 6-merous flowers can also be seen. Most <italic>Rhoiacarpos</italic> flowers are 5-merous, but 4- and 6-merous flowers are also known. The number of lobes of the glandular disk in these genera follows the number of petals.</p>
</sec>
<sec id="s20007">
<title>Fruits</title>
<p>In all three genera, many fewer fruits develop to maturity compared with the number of flowers originally present on the inflorescence axis. In <italic>Colpoon</italic>, evidence of flower abortion exists (<xref ref-type="fig" rid="F0003">Figure 3d</xref>) such that a single fruit is present per axis with the scars of abortive flowers present at the swollen peduncle apex (<xref ref-type="fig" rid="F0003">Figure 3f</xref>). The fruits of <italic>Rhoiacarpos</italic> differ from the other two genera in having persistent petals at the apex (<xref ref-type="fig" rid="F0004">Figure 4f</xref> and <xref ref-type="fig" rid="F0004">g</xref>). A cluster of fruits is shown in <xref ref-type="fig" rid="F0004">Figure 4f</xref>; however, each fruit appears to be present on a separate inflorescence branch with evidence of numerous abortive flowers and their associated bracts below (<xref ref-type="fig" rid="F0004">Figure 4g</xref>).</p>
<p>In <italic>Osyris</italic> (<xref ref-type="fig" rid="F0002">Figure 2f</xref>) and <italic>C. compressum</italic> (<xref ref-type="fig" rid="F0003">Figure 3f</xref>), the petals abscise and are not persistent on the mature fruit. Bean (<xref ref-type="bibr" rid="CIT0003">1990</xref>) indicates that in <italic>C. speciosum</italic> the petals are persistent; thus, this feature is polymorphic within the genus. Although it is not clear whether the character of fruit shape will remain valid following further observations, it appears that the fruit in <italic>Colpoon</italic> is more obovoid than in the other two genera that have spheroid to ellipsoid fruits.</p>
</sec>
</sec>
<sec id="s0008">
<title>Discussion</title>
<p>The decision by Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>) to lump <italic>Colpoon</italic> into <italic>Osyris</italic> can now be addressed given the molecular phylogenetic and comparative morphology results. From the topologies of the phylogenetic trees (<xref ref-type="fig" rid="F0001">Figure 1a</xref> and <xref ref-type="fig" rid="F0001">b</xref>), one could include <italic>Colpoon</italic> in <italic>Osyris</italic>. If this was done, and monophyly was maintained, then the North American <italic>Nestronia</italic> and South African <italic>Rhoiacarpos</italic> would also have to be included in <italic>Osyris</italic>. This concept has never been proposed and in fact <italic>Rhoiacarpos</italic> has been considered a separate genus in all consulted works on South African flora. Molecular dating conducted by Vidal-Russell and Nickrent (<xref ref-type="bibr" rid="CIT0014">2008</xref>) showed that <italic>Santalum</italic> and <italic>Osyris</italic> diverged in the Paleocene, at least 60 million years ago. Although the other genera (<italic>Nestronia, Rhoiacarpos</italic> and <italic>Colpoon</italic>) have not been included in ultrametric trees, it is likely that they diverged from each other somewhat more recently, for example, in the Eocene, ca. 50 million years ago.</p>
<p>Despite the statement by Hill (<xref ref-type="bibr" rid="CIT0007">1915</xref>) that &#x2018;There is also no floral difference between <italic>Colpoon</italic> Berg. and <italic>Osyris</italic> Linn&#x2019;. and the similar sentiment expressed by Hilliard (<xref ref-type="bibr" rid="CIT0008">1994</xref>), the evidence presented above shows that there are clear differences in vegetative, floral and fruit features among all three genera. The taxonomic conclusions reached by Stauffer (<xref ref-type="bibr" rid="CIT0012">1961</xref>) are fully supported by molecular phylogenetic data; thus, <italic>Colpoon</italic> should be considered a genus, distinct from <italic>Osyris</italic>, and is composed of two species, <italic>C. compressum</italic> and <italic>C. speciosum</italic>.</p>
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<ack>
<title>Acknowledgements</title>
<p>I thank Marinda Koekemoer for allowing me to use several photos of <italic>Osyris lanceolata</italic> and <italic>Colpoon compressum</italic>. Photos of <italic>Rhoiacarpos capensis</italic>, posted online at <ext-link ext-link-type="uri" xlink:href="http://www.floraoftheworld.org">www.floraoftheworld.org</ext-link>, were taken by Christopher Davidson. One of the reviewers is acknowledged for pointing out overlooked but important literature.</p>
<sec id="s20009" sec-type="COI-statement">
<title>Competing interests</title>
<p>The author declares that he has no financial or personal relationships that may have inappropriately influenced him in writing this article.</p>
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<fn-group>
<fn><p><bold>How to cite this article:</bold> Nickrent, D.L, 2017, &#x2018;Status of the Genera <italic>Colpoon, Osyris</italic> and <italic>Rhoiacarpos</italic> in South Africa&#x2019;, <italic>Bothalia</italic> 47(1), a2260. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.4102/abc.v47i1.2260">https://doi.org/10.4102/abc.v47i1.2260</ext-link></p></fn>
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