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<document ID-DOI="http://dx.doi.org/10.3897/phytokeys.85.14757" ID-PMC="PMC5624194" ID-Pensoft-Pub="1314-2003-85-69" ID-Pensoft-UUID="FFE8FF963B157F1D747CFFF4FFBDFFB4" ID-PubMed="29033661" ID-Zenodo-Dep="899159" ModsDocID="1314-2003-85-69" checkinTime="1504194347456" checkinUser="pensoft" docAuthor="Wurdack, Kenneth J. &amp; Farfan-Rios, William" docDate="2017" docId="4935F12549BE542DB00233FC564840D3" docLanguage="en" docName="PhytoKeys 85: 69-86" docOrigin="PhytoKeys 85" docPubDate="2017-08-31" docSource="http://dx.doi.org/10.3897/phytokeys.85.14757" docTitle="Incadendron esseri K. Wurdack &amp; Farfan 2017, sp. nov." docType="treatment" docVersion="3" id="FFE8FF963B157F1D747CFFF4FFBDFFB4" lastPageNumber="81" masterDocId="FFE8FF963B157F1D747CFFF4FFBDFFB4" masterDocTitle="Incadendron: a new genus of Euphorbiaceae tribe Hippomaneae from the sub-Andean cordilleras of Ecuador and Peru" masterLastPageNumber="86" masterPageNumber="69" pageNumber="71" updateTime="1668142046557" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Incadendron: a new genus of Euphorbiaceae tribe Hippomaneae from the sub-Andean cordilleras of Ecuador and Peru</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Wurdack, Kenneth J.</mods:namePart>
<mods:affiliation>Department of Botany, MRC- 166, National Museum of Natural History, Smithsonian Institution, P. O. Box 37012, Washington DC 20013 - 7012, USA</mods:affiliation>
<mods:nameIdentifier type="email">wurdackk@si.edu</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Farfan-Rios, William</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-3196-0317</mods:nameIdentifier>
<mods:affiliation>Department of Biology, Wake Forest University, Winston-Salem, NC 27106, USA</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>PhytoKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2017</mods:date>
<mods:detail type="pubDate">
<mods:number>2017-08-31</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>85</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>69</mods:start>
<mods:end>86</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/phytokeys.85.14757</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/phytokeys.85.14757</mods:identifier>
<mods:identifier type="Pensoft-Pub">1314-2003-85-69</mods:identifier>
<mods:identifier type="Pensoft-UUID">FFE8FF963B157F1D747CFFF4FFBDFFB4</mods:identifier>
<mods:identifier type="Zenodo-Dep">899159</mods:identifier>
</mods:mods>
<subSection lastPageId="12" lastPageNumber="81" pageId="2" pageNumber="71" type="multiple">
<treatment ID-GBIF-Taxon="182225816" LSID="urn:lsid:plazi:treatment:4935F12549BE542DB00233FC564840D3" httpUri="http://treatment.plazi.org/id/4935F12549BE542DB00233FC564840D3" lastPageId="12" lastPageNumber="81" pageId="2" pageNumber="71">
<subSubSection pageId="2" pageNumber="71" type="nomenclature">
<paragraph pageId="2" pageNumber="71">
<taxonomicName LSID="4935F125-49BE-542D-B002-33FC564840D3" authority="K. Wurdack &amp; Farfan" authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron esseri" order="Malpighiales" pageId="2" pageNumber="71" phylum="Tracheophyta" rank="species" species="esseri" status="sp. nov.">Incadendron esseri K.Wurdack &amp; Farfan</taxonomicName>
<taxonomicNameLabel pageId="2" pageNumber="71">sp. nov.</taxonomicNameLabel>
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Illustration of Incadendron esseri. A Habit B Shoot tip C Leaf base (adaxial) D Leaf base and marginal glands (abaxial). E Staminate subinflorescence F Staminate cymule (distal view) G Staminate cymule (proximal view, without lateral buds) H Staminate flower I Pistillate flower J Fruit K Mericarp valve L Columella M Seed (ventral face). (Source: A-G Calatayud et al. 4711, MO; H-I Monteagudo et al. 4458, US; J Vasquez &amp; Valenzuela 37638, MO; K-M Monteagudo &amp; Ortiz 4605, US)." figureDoi="10.3897/phytokeys.85.14757.figure1" httpUri="https://binary.pensoft.net/fig/153291" pageId="2" pageNumber="71">Figure 1</figureCitation>
</paragraph>
</subSubSection>
<subSubSection pageId="2" pageNumber="71" type="materials_examined">
<paragraph pageId="2" pageNumber="71">
<typeStatus>Type</typeStatus>
.
</paragraph>
<paragraph pageId="2" pageNumber="71">
<materialsCitation collectingDate="2007-09-24" collectorName="La Convencion, de Yavero, G. Calatayud, I. Huamantupa, E. Suclli, R. Ayerbe" country="PERU" county="La Convencion" elevation="2301" latitude="-12.478611" location="Abra de Yavero" longLatPrecision="21" longitude="-72.48333" municipality="Districto Quellouno" specimenCount="1" stateProvince="Cusco" typeStatus="holotype">
<collectingCountry name="Peru">PERU</collectingCountry>
.
<collectingRegion country="Peru" name="Cusco">Cusco</collectingRegion>
:
<collectorName>
<collectingCounty>
La
<normalizedToken originalValue="Convención">Convencion</normalizedToken>
</collectingCounty>
</collectorName>
,
<collectingMunicipality>Districto Quellouno</collectingMunicipality>
,
<location LSID="urn:lsid:plazi:treatment:4935F12549BE542DB00233FC564840D3:176A087F068AF51C9E5ED0C8FA585A3D" country="PERU" county="La Convencion" latitude="-12.478611" longLatPrecision="21" longitude="-72.48333" municipality="Districto Quellouno" name="Abra de Yavero" stateProvince="Cusco">
Abra
<collectorName>de Yavero</collectorName>
</location>
,
<geoCoordinate degrees="12" direction="south" minutes="28" orientation="latitude" precision="15" seconds="43" value="-12.478611">12°28'43&quot;S</geoCoordinate>
<geoCoordinate degrees="072" direction="west" minutes="29" orientation="longitude" precision="15" seconds="00" value="-72.48333">072°29'00&quot;W</geoCoordinate>
,
<quantity metricMagnitude="3" metricUnit="m" metricValue="2.3009999999999997" unit="m" value="2301.0">
<elevation metricMagnitude="3" metricUnit="m" metricValue="2.3009999999999997" unit="m" value="2301.0">2301 m</elevation>
</quantity>
,
<collectingDate value="2007-09-24">24 Sep 2007</collectingDate>
(fl, fr green fide label),
<emphasis italics="true" pageId="2" pageNumber="71">
<collectorName>G. Calatayud</collectorName>
,
<collectorName>I. Huamantupa</collectorName>
,
<collectorName>E. Suclli</collectorName>
, &amp;
<collectorName>R. Ayerbe</collectorName>
4711
</emphasis>
(
<typeStatus>holotype</typeStatus>
: USM; isotypes: AMAZ, CUZ, HUT, MO-6669029, MOL, US-3679263)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="4" lastPageNumber="73" pageId="2" pageNumber="71" type="description">
<paragraph pageId="2" pageNumber="71">Description.</paragraph>
<paragraph lastPageId="4" lastPageNumber="73" pageId="2" pageNumber="71">
<emphasis italics="true" pageId="2" pageNumber="71">Trees</emphasis>
6-26 m tall, to at least 56.7 cm dbh, trunk bark thin, monoecious; flowering and fruiting branchlets with leaves present, branching in pairs, leafy stems 1.5-2 mm dia., internodes 1-1.5 cm apart, branchlet bark smooth with scattered leaf and stipule scars, pith soft.
<emphasis italics="true" pageId="2" pageNumber="71">Exudate</emphasis>
present, white latex, watery.
<emphasis italics="true" pageId="2" pageNumber="71">Indument</emphasis>
absent.
<emphasis italics="true" pageId="2" pageNumber="71">Leaves</emphasis>
alternate, petiolate, stipulate, simple.
<emphasis italics="true" pageId="2" pageNumber="71">Stipules</emphasis>
free, paired, overlapping to sheath terminal bud, lanceolate, 10-13
<normalizedToken originalValue="×">x</normalizedToken>
3-4 mm, tip acute; base cordate, slightly asymmetric, with rounded free lobes extending to 1 mm below central point of attachment; eglandular, margins hyaline, deciduous before new leaf has fully expanded (abscised before leaf is 1/3 of mature size), after abscission leaving elliptic to reniform scars 1.3-1.7
<normalizedToken originalValue="×">x</normalizedToken>
0.4-0.5 mm.
<emphasis italics="true" pageId="2" pageNumber="71">Petioles</emphasis>
9-20
<normalizedToken originalValue="×">x</normalizedToken>
1-1.8 mm (dia. mid-length), slightly flared at base, adaxially canaliculate, groove shallow and wide at petiole base (proximal) then narrowed mid-length and finally deep and wide at distal apex where shoulders of grove support minutely auriculate leaf base, petiolar glands absent.
<emphasis italics="true" pageId="2" pageNumber="71">Leaf blades</emphasis>
: laminar size class microphyll to notophyll, blade 6.7-13
<normalizedToken originalValue="×">x</normalizedToken>
2.9-3.8 cm, length:width ratio 1.97-
<pageBreakToken pageId="3" pageNumber="72" start="start">2.89</pageBreakToken>
:1 (mean = 2.49, SD = 0.212, n = 40, 2-3 mature leaves each from 14 collections), symmetrical, elliptic to slightly obovate, apex acute to rounded, often minutely retuse, base acute and minutely auriculate at point of attachment; margin with distinct smooth edge ca 0.1 mm wide, slightly revolute with more pronounced inward rolling at leaf base; marginal glands up to 12 per side with 2-4 larger ones consistently near base then sparse and progressively smaller distally, glands embedded in leaf margin (those at base hidden or appearing abaxial due to rolling of margin) and often appear perpendicular to blade, 0.2-1
<normalizedToken originalValue="×">x</normalizedToken>
0.1-0.5 mm, narrowly elliptic, surface slightly sunken, smooth; apex of midvein (where terminating at margin zone) minutely apiculate or thickened with globular mass 0.2 mm dia.; lamina coriaceous, adaxial surface of new leaves glossy, abaxially appearing smooth; ptyxis conduplicate, halves of young blade folded tightly together along adaxial surface; embedded laminar glands absent.
<emphasis italics="true" pageId="3" pageNumber="72">Venation</emphasis>
pinnate, brochidodromous; secondaries (10)13-17(19) pairs, spacing uniform, vein angle uniform to decreasing proximally, (10)20-30(40)°, decurrent attachment to midvein; intersecondaries frequent, parallel to secondaries; intercostal tertiaries reticulate; primary to tertiary venation prominulous on both surfaces.
<emphasis italics="true" pageId="3" pageNumber="72">Inflorescence</emphasis>
bisexual or staminate, solitary, terminal but appearing leaf-opposed, spicate thyrse, 5-12 cm long including 0.9-1.8 cm peduncle and distal rachis; in young bud protected by adjacent sheathing leaf stipules and bracts; bracts 2, free to shortly connate, stipuliform, inserted at start of fertile part (i.e., just below pistillate flower), 7
<normalizedToken originalValue="×">x</normalizedToken>
3.5 mm, rarely with 1-2 glands at base, cauducous.
<emphasis italics="true" pageId="3" pageNumber="72">Flowers</emphasis>
unisexual, lacking petals, disc, staminodes, or pistillodes.
<emphasis italics="true" pageId="3" pageNumber="72">Staminate partial inflorescence</emphasis>
a lax spiral of numerous cymules, each cymule on a tissue pad bearing a bract and subtended by glands; cymule bracts ca 0.5 (free portion)
<normalizedToken originalValue="×">x</normalizedToken>
1 mm, widely acute to rounded, verruculose, persistent, when young forming protective scale-like sheath around staminate subinflorescence and inclinate buds; glands 4-5(6) in row or cluster per side of bract, 0.5-0.6
<normalizedToken originalValue="×">x</normalizedToken>
0.3-0.4 mm, disc-shaped or prismoidal when tightly abutting each other, face concave, yellow-orange in life; flowers 3 per cymule, central flower precocious and senescent well before laterals; bracteoles absent.
<emphasis italics="true" pageId="3" pageNumber="72">Staminate flowers</emphasis>
erect at anthesis; pedicels 3
<normalizedToken originalValue="×">x</normalizedToken>
0.3 mm for central flower (laterals seen only in bud), articulated at base; sepals 3, connate to 0.2 mm, widely rounded, 0.5
<normalizedToken originalValue="×">x</normalizedToken>
0.7 mm; stamens 3, free; filaments distinct, shorter than anthers, to 0.3 mm long; anthers 0.6
<normalizedToken originalValue="×">x</normalizedToken>
0.6 mm, 2-thecate, basifixed, exserted slightly through sinus between sepal lobes, dehiscent through longitudinal slits to 0.3 mm long, slit margins slightly recurved at anthesis; connective tip acute, barely protruding beyond thecae, not elaborated; yellow in life.
<emphasis italics="true" pageId="3" pageNumber="72">Pollen</emphasis>
polar:equatorial ratio (1.13)1.16-1.25:1 (based on SEM), tectum perforations smaller near apertures.
<emphasis italics="true" pageId="3" pageNumber="72">Pistillate flower</emphasis>
: (0)1, basal, pedicellate; pedicel 3.5-4.5
<normalizedToken originalValue="×">x</normalizedToken>
0.8-1 mm; pistillate bracts (1)2, at base of pedicel (shortly distal to stipuliform inflorescence bracts), 1-4
<normalizedToken originalValue="×">x</normalizedToken>
0.6-1 mm, elliptic to lanceolate, cauducous, leaving scars 0.3
<normalizedToken originalValue="×">x</normalizedToken>
0.2 mm; glands absent; flower 6-9 mm long; sepals 3, connate to 1 mm, broadly acute, 1.7-3.5
<normalizedToken originalValue="×">x</normalizedToken>
1.8-1.9 mm, margins hyaline; ovary 3-locular, ca 2
<normalizedToken originalValue="×">x</normalizedToken>
2 mm, top tapering and barely distinguished from start of styles; placentation apical pendulous with single ovule per locule; styles connate into column ca 5
<normalizedToken originalValue="×">x</normalizedToken>
0.9-1 mm; stigmas 3, free, undivided, eglandular, ca 3 mm long, erect in bud, recurved to loosely coiled at anthesis, surface
<pageBreakToken pageId="4" pageNumber="73" start="start">minutely</pageBreakToken>
papillose.
<emphasis italics="true" pageId="4" pageNumber="73">Infructescence</emphasis>
axis 23-40 mm long, consisting of peduncle 11-20
<normalizedToken originalValue="×">x</normalizedToken>
1.3-1.5 mm (dia. mid-length) and pistillate flower pedicel 10-20
<normalizedToken originalValue="×">x</normalizedToken>
1-1.3 mm (dia. mid-length and distinctly thinner than peduncle), prominent scars where staminate partial inflorescence and bracts abscised.
<emphasis italics="true" pageId="4" pageNumber="73">Fruit</emphasis>
subglobose, trigonous, 20
<normalizedToken originalValue="×">x</normalizedToken>
20 mm; ventral (septal) sutures sulcate; dorsal (loculicidal) sutures smooth when fresh, becoming ridged when dry; apex with woody beak ca 3 mm tall, sepals and stigmas deciduous; mericarps equal, 2-valved, splitting septicidally then loculicidally to release seeds.
<emphasis italics="true" pageId="4" pageNumber="73">Pericarp</emphasis>
3-4 mm thick (equatorial at dorsal suture); dried exocarp thin, 0.2 mm mid-mericarp to 1 mm at ventral septal sutures and 2 mm at base, loosely adherent (can be easily peeled off dry specimens) to woody mesocarp on dehiscence, prominulous (likely drying artifact) veined with one primary vein descending from apex per valve, lower vein orders reticulate, major venation tracking embossed ridges on woody mesocarp; mericarp valves (cocci) barely twisted when dehisced, remaining attached together via basal triangle 3-5
<normalizedToken originalValue="×">x</normalizedToken>
6-7 mm (width at base); septa of mericarps woody, nearly continuous except for small semi-circular gap 2 (wide)
<normalizedToken originalValue="×">x</normalizedToken>
1 mm (deep) where traversed by funicle; abscission layer between septa with well-developed spongy layer except absent at beak, vascular strand single or bifurcate; columella (carpophore) 18-20 mm long, proximally (where traversing pericarp at fruit base) rounded, distally (where confluent with septa) trigonous, distal part alate with jagged wing extending to 2 mm from central axis; funicle erect, slender, 2
<normalizedToken originalValue="×">x</normalizedToken>
0.3 mm.
<emphasis italics="true" pageId="4" pageNumber="73">Seeds</emphasis>
ellipsoid, apically (hilar end) rounded, basally flattened to slightly depressed, 8.8-9.7 (long)
<normalizedToken originalValue="×">x</normalizedToken>
6.6-7.5 (wide; lateral-lateral)
<normalizedToken originalValue="×">x</normalizedToken>
6.6-7 mm (deep; raphe-antiraphe), ventral face longitudinally traversed by narrow raised raphe 0.2 mm wide; seed coat ca 0.1 mm thick, testa thin, dry, uniformly dark brown; exotegmen mechanical, uniseriate palisade layer of elongate thick-walled cells, cells 3-4
<normalizedToken originalValue="×">x</normalizedToken>
shorter at apex versus bottom and sides, cell orientation vertical, inclined, or curved depending on location; caruncle absent; seed contents separated from mechanical coat by thin spongy layer except at base where solidly attached; endosperm yellowish, fleshy, slightly oily, forming layer up to 1 mm thick around central flattened ellipsoidal pocket containing embryo with cotyledons adhering to side of pocket; embryo type spathulate fully developed, embryo straight, extending most of seed length; cotyledons flat, elliptic, 4.5
<normalizedToken originalValue="×">x</normalizedToken>
3 mm, thin (ca 0.1 mm), apex broadly rounded, base subcordate, prominulous central vein that is distally branched; hypocotyl-radicle (stalk sensu
<bibRefCitation DOI="https://doi.org/10.2307/2421457" author="Martin, AC" journalOrPublisher="The American Midland Naturalist" pageId="14" pageNumber="83" pagination="513 - 660" refId="B16" refString="Martin, AC, 1946. The comparative internal morphology of seeds. The American Midland Naturalist 36: 513 - 660, DOI: https://doi.org/10.2307/2421457" title="The comparative internal morphology of seeds." url="https://doi.org/10.2307/2421457" volume="36" year="1946">Martin 1946</bibRefCitation>
) 2
<normalizedToken originalValue="×">x</normalizedToken>
1 mm, laterally slightly flattened.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="73" type="etymology">
<paragraph pageId="4" pageNumber="73">Etymology.</paragraph>
<paragraph pageId="4" pageNumber="73">
The genus is combined from &quot;
<emphasis italics="true" pageId="4" pageNumber="73">Inca</emphasis>
&quot; (as
<emphasis italics="true" pageId="4" pageNumber="73">Inka</emphasis>
, Quechua for
<normalizedToken originalValue="“ruler”">&quot;ruler&quot;</normalizedToken>
or
<normalizedToken originalValue="“lord”">&quot;lord&quot;</normalizedToken>
) referring to the indigenous Inca people and pre-Columbian empire that was centered in Cusco and encompassed much of the range of this taxon, and &quot;
<emphasis italics="true" pageId="4" pageNumber="73">dendron</emphasis>
&quot; (Greek) referring to tree, which is the habit of the plant. Some localities occur near the Trocha
<normalizedToken originalValue="Unión">Union</normalizedToken>
, an ancient Inca path. The specific epithet is from &quot;
<emphasis italics="true" pageId="4" pageNumber="73">Esser</emphasis>
&quot;, the surname of Hans-Joachim Esser (Botanische Staatssammlung
<normalizedToken originalValue="München">Muenchen</normalizedToken>
, Germany) and honors this expert on Hippomaneae who has contributed much to our understanding of the tribe and
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" family="Euphorbiaceae" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="4" pageNumber="73" rank="family">Euphorbiaceae</taxonomicName>
in general.
</paragraph>
</subSubSection>
<subSubSection lastPageId="6" lastPageNumber="75" pageId="4" pageNumber="73" type="distribution">
<paragraph pageId="4" pageNumber="73">Distribution, life history, and ecology.</paragraph>
<paragraph lastPageId="6" lastPageNumber="75" pageId="4" pageNumber="73">
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="4" pageNumber="73" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="4" pageNumber="73">Incadendron</emphasis>
</taxonomicName>
is known from three well-separated clusters of localities (hereafter referred to as
<normalizedToken originalValue="Cóndor">Condor</normalizedToken>
, Manu, and Oxapampa
<pageBreakToken pageId="5" pageNumber="74" start="start">populations</pageBreakToken>
) on the eastern slopes of the main Andean mountain range in Peru and Ecuador, where it occurs in wet montane forests at 1800-2400 m elevation (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Distribution map of Incadendron esseri." figureDoi="10.3897/phytokeys.85.14757.figure2" httpUri="https://binary.pensoft.net/fig/153292" pageId="5" pageNumber="74">2</figureCitation>
). The extent of discontinuity in its range is presently unclear due to the floristically poorly known nature of the intervening areas, and it should be looked for in similar habitats between the three populations. There are minor vegetative differences including leaf apex variation with most tips distinctly acute versus more rarely rounded (i.e.,
<emphasis italics="true" pageId="6" pageNumber="75">
<pageBreakToken pageId="6" pageNumber="75" start="start">Neill</pageBreakToken>
&amp; Kajekai 16620
</emphasis>
,
<emphasis italics="true" pageId="6" pageNumber="75">Monteagudo et al. 16929</emphasis>
), and a larger-leafed collection (i.e.,
<emphasis italics="true" pageId="6" pageNumber="75">Monteagudo et al. 4458</emphasis>
). The differences exist within the populations and presently do not suggest differentiation worthy of taxonomic recognition.
</paragraph>
<paragraph pageId="6" pageNumber="75">
Detailed field observations were made in the tropical montane cloud forests of the
<normalizedToken originalValue="Kosñipata">Kosnipata</normalizedToken>
Valley in Manu National Park (Parque Nacional del Manu), which contains the southernmost part of the known range of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Incadendron</emphasis>
</taxonomicName>
. The general site characteristics are well documented (see
<bibRefCitation DOI="https://doi.org/10.1111/j.1365-2486.2010.02235.x" author="Girardin, CA" journalOrPublisher="Global Change Biology" pageId="14" pageNumber="83" pagination="3176 - 3192" refId="B13" refString="Girardin, CA, Malhi, Y, Aragao, LEOC, Mamani, M, Huaraca, Huasco W, Durand, L, Feely, KJ, Rapp, J, Silva-Espejo, JE, Silman, MR, Salinas, N, Whittaker, RJ, 2010. Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes. Global Change Biology 16: 3176 - 3192, DOI: https://doi.org/10.1111/j.1365-2486.2010.02235.x" title="Net primary productivity allocation and cycling of carbon along a tropical forest elevational transect in the Peruvian Andes." url="https://doi.org/10.1111/j.1365-2486.2010.02235.x" volume="16" year="2010">Girardin et al. 2010</bibRefCitation>
,
<bibRefCitation author="Farfan-Rios, W" journalOrPublisher="MSc Thesis, Wake Forest University, USA" pageId="13" pageNumber="82" refId="B10" refString="Farfan-Rios, W, 2011. Changes in forest dynamics along a 2.5 km elevation gradient on the southeastern flank of the Peruvian Andes. MSc Thesis, Wake Forest University, USA" title="Changes in forest dynamics along a 2.5 km elevation gradient on the southeastern flank of the Peruvian Andes." year="2011">Farfan Rios 2011</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.1111/j.1365-2699.2010.02444.x" author="Feeley, KJ" journalOrPublisher="Journal of Biogeography" pageId="14" pageNumber="83" pagination="783 - 791" refId="B12" refString="Feeley, KJ, Silman, MR, Bush, MB, Farfan, W, Cabrera, KG, Malhi, Y, Saatchi, S, 2011. Upslope migration of Andean trees. Journal of Biogeography 38: 783 - 791, DOI: https://doi.org/10.1111/j.1365-2699.2010.02444.x" title="Upslope migration of Andean trees." url="https://doi.org/10.1111/j.1365-2699.2010.02444.x" volume="38" year="2011">Feeley et al. 2011</bibRefCitation>
, Rapp et al. 2012) as part of intensive forest monitoring using permanent forests plots established by the Andes Biodiversity and Ecosystem Research Group (ABERG, http://www.andesconservation.org/) along an elevational gradient (i.e.,
<normalizedToken originalValue="Kosñipata">Kosnipata</normalizedToken>
transect) from the Andes to the Amazon.
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Incadendron</emphasis>
</taxonomicName>
has been found (i.e.,
<emphasis italics="true" pageId="6" pageNumber="75">Farfan et al. 1049</emphasis>
,
<emphasis italics="true" pageId="6" pageNumber="75">1090</emphasis>
,
<emphasis italics="true" pageId="6" pageNumber="75">1131</emphasis>
) in the cloud immersion zone between 1800-2250 m at the study site. The substrate where the tree was collected is granite between 1800-2000 m, and shale at 2250 m. The soils below the thick organic surface layer are relatively poor in nutrients. At the elevations where found,
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Incadendron</emphasis>
</taxonomicName>
is among the taller components of the forest and its habit is a canopy tree with a spreading crown. The maximum height observed was 21.2 m, and maximum tree diameter at breast height (dbh) was 56.7 cm. When cut, the thin trunk bark has a cream-yellowish color with abundant white latex. Mean tree growth (diameter increment) at the study site was 4.02 mm yr-1
<normalizedToken originalValue="±">+/-</normalizedToken>
0.90 (95% CI). Mean wood density is 0.55 g/cm3
<normalizedToken originalValue="±">+/-</normalizedToken>
1.18 (95% CI), based in field sampling. The highest population density in the network of one hectare plots was found at 2000 m of elevation, with 30 adult individuals/ha (≥10 cm dbh), making it the ninth most abundant tree in that plot (Parcela VII). The main associated species include
<taxonomicName class="Magnoliopsida" family="Alzateaceae" genus="Alzatea" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Alzatea verticillata" order="Myrtales" pageId="6" pageNumber="75" phylum="Magnoliophyta" rank="species" species="verticillata">
<emphasis italics="true" pageId="6" pageNumber="75">Alzatea verticillata</emphasis>
</taxonomicName>
Ruiz &amp; Pav. (
<taxonomicName authorityName="Ruiz &amp; Pavon" authorityYear="1794" class="Magnoliopsida" family="Alzateaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Myrtales" pageId="6" pageNumber="75" phylum="Magnoliophyta" rank="family">Alzateaceae</taxonomicName>
),
<taxonomicName authorityName="Mett" authorityYear="1859" class="Filicopsida" family="Cyatheaceae" genus="Cyathea" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Cyathea lechleri" order="Cyatheales" pageId="6" pageNumber="75" phylum="Filicinophyta" rank="species" species="lechleri">
<emphasis italics="true" pageId="6" pageNumber="75">Cyathea lechleri</emphasis>
</taxonomicName>
Mett. (
<taxonomicName class="Filicopsida" family="Cyatheaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Cyatheales" pageId="6" pageNumber="75" phylum="Filicinophyta" rank="family">Cyatheaceae</taxonomicName>
), and
<taxonomicName authorityName="Loes" authorityYear="1901" class="Magnoliopsida" family="Euphorbiaceae" genus="Ilex" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Ilex villosula" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="species" species="villosula">
<emphasis italics="true" pageId="6" pageNumber="75">Ilex villosula</emphasis>
</taxonomicName>
Loes. (
<taxonomicName class="Dicotyledoneae" family="Aquifoliaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Celastrales" pageId="6" pageNumber="75" phylum="Angiospermae" rank="family">Aquifoliaceae</taxonomicName>
). The
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="family">Euphorbiaceae</taxonomicName>
diversity for the tropical montane forests of this region includes nine genera, of which there is notable species-richness in
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Alchornea" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Alchornea" order="Malpighiales" pageId="6" pageNumber="75" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Alchornea</emphasis>
</taxonomicName>
Sw. (
<bibRefCitation DOI="https://doi.org/10.15381/rpb.v22i2.11351" author="Farfan-Rios, W" journalOrPublisher="Revista Peruana de Biologia" pageId="13" pageNumber="82" pagination="145 - 174" refId="B11" refString="Farfan-Rios, W, Garcia-Cabrera, K, Salinas, N, Silman, MR, 2015. Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando. Revista Peruana de Biologia 22: 145 - 174, DOI: https://doi.org/10.15381/rpb.v22i2.11351" title="Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando." url="https://doi.org/10.15381/rpb.v22i2.11351" volume="22" year="2015">Farfan-Rios et al. 2015</bibRefCitation>
). The closet relatives of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Incadendron</emphasis>
</taxonomicName>
(i.e., other members of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" family="Euphorbiaceae" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="6" pageNumber="75" rank="subFamily" subFamily="Euphorbioideae">Euphorbioideae</taxonomicName>
) among these nine genera include
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sapium" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Sapium" order="Malpighiales" pageId="6" pageNumber="75" phylum="Angiospermae" rank="genus">
<emphasis italics="true" pageId="6" pageNumber="75">Sapium</emphasis>
</taxonomicName>
spp. and
<taxonomicName authorityName="Esser" authorityYear="2001" baseAuthorityName="Müll. Arg." class="Magnoliopsida" family="Euphorbiaceae" genus="Pseudosenefeldera" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Pseudosenefeldera inclinata" order="Lepidoptera" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="species" species="inclinata">
<emphasis italics="true" pageId="6" pageNumber="75">Pseudosenefeldera inclinata</emphasis>
</taxonomicName>
(
<normalizedToken originalValue="Müll">Muell</normalizedToken>
. Arg.) Esser, with the latter occurring at lower elevations. The basal marginal leaf glands of
<emphasis italics="true" pageId="6" pageNumber="75">Neill &amp; Kajekai 16620</emphasis>
from the Cordillera del
<normalizedToken originalValue="Cóndor">Condor</normalizedToken>
are overgrown by unusual clusters of tiny, 0.1 mm diameter black globules that are fruiting bodies of a likely ascomycete fungus. While epiphyllous fungal growth such as mold growing on glandular secretions is to be expected, these unusual fruiting structures are very different and deserve further study.
</paragraph>
</subSubSection>
<subSubSection lastPageId="7" lastPageNumber="76" pageId="6" pageNumber="75" type="phenology">
<paragraph pageId="6" pageNumber="75">Phenology.</paragraph>
<paragraph lastPageId="7" lastPageNumber="76" pageId="6" pageNumber="75">
The trees are evergreen, with an observed flowering season during July-September and fruiting during August-November. Herbarium collections also indicate a spring reproductive period of January-April for the Manu and Oxapampa populations. Fruits can be abundant, they turn brown when mature (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="6" pageNumber="75">3G-H</figureCitation>
), and due to their large, heavy nature become pendulous on the relatively long infructescence axes. They are subject to predispersal seed predation by
<taxonomicName authorityName="H.-J.Esser" authorityYear="2001" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Lepidoptera" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="order">Lepidoptera</taxonomicName>
, based on caterpillar remains recovered from inside fragmented fruits of
<emphasis italics="true" pageId="6" pageNumber="75">Monteagudo &amp; Ortiz 4605</emphasis>
. In developing fruits these moths (likely members of
<taxonomicName lsidName="" pageId="6" pageNumber="75" rank="subFamily" subFamily="Phycitinae">Phycitinae</taxonomicName>
,
<taxonomicName class="Insecta" family="Pyralidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" lsidName="" order="Lepidoptera" pageId="6" pageNumber="75" phylum="Arthropoda" rank="family">Pyralidae</taxonomicName>
:
<taxonomicName authorityName="H.-J.Esser" authorityYear="2001" class="Magnoliopsida" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Lepidoptera" pageId="6" pageNumber="75" phylum="Tracheophyta" rank="superFamily" superFamily="Pyraloidea">Pyraloidea</taxonomicName>
; A. Solis, personal communication) hollow out the seeds, which have well-developed endosperm, and leave holes (1.75 mm dia.) in the mericarp septa and seed coats (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="6" pageNumber="75">3I-J</figureCitation>
). Plant
<pageBreakToken pageId="7" pageNumber="76" start="start">defenses</pageBreakToken>
to deter herbivory would appear to be high in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="7" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="7" pageNumber="76">Incadendron</emphasis>
</taxonomicName>
due to latex, and the thick, lignified pericarp. Seed predation by specialist moths is well known for other Hippomaneae including in
<taxonomicName authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Euphorbiaceae" genus="Mabea" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Mabea" order="Malpighiales" pageId="7" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="7" pageNumber="76">Mabea</emphasis>
</taxonomicName>
Aubl., where oviposition occurs early in fruit development when the pericarp is thin and soft (
<bibRefCitation DOI="https://doi.org/10.2307/2387717" author="De Steven, D" journalOrPublisher="Biotropica" pageId="13" pageNumber="82" pagination="146 - 150" refId="B5" refString="De Steven, D, 1981. Predispersal seed predation in a tropical shrub (Mabea occidentialis, Euphorbiaceae). Biotropica 13: 146 - 150, DOI: https://doi.org/10.2307/2387717" title="Predispersal seed predation in a tropical shrub (Mabea occidentialis, Euphorbiaceae)." url="https://doi.org/10.2307/2387717" volume="13" year="1981">De Steven 1981</bibRefCitation>
). One young
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="7" pageNumber="76" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="7" pageNumber="76">Incadendron</emphasis>
</taxonomicName>
fruit (4 mm dia.) on
<emphasis italics="true" pageId="7" pageNumber="76">Monteagudo &amp; Ortiz 4605</emphasis>
(US) has what appears to be an oviposition hole at the top of the ovary that is likely a weak spot into the interior.
</paragraph>
</subSubSection>
<subSubSection pageId="7" pageNumber="76" type="conservation status">
<paragraph pageId="7" pageNumber="76">Conservation status.</paragraph>
<paragraph pageId="7" pageNumber="76">
Following the criteria and categories of
<bibRefCitation author="IUCN" journalOrPublisher="Global Change Biology" pageId="14" pageNumber="83" refId="B14" refString="IUCN, 2012. IUCN Red List Categories and Criteria: Version 3.1. Second edition. IUCN, Gland &amp; Cambridge, iv + 32 pp." title="IUCN Red List Categories and Criteria: Version 3.1. Second edition. IUCN, Gland &amp; Cambridge, iv + 32 pp." year="2012">IUCN (2012)</bibRefCitation>
,
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron esseri" order="Malpighiales" pageId="7" pageNumber="76" phylum="Tracheophyta" rank="species" species="esseri">
<emphasis italics="true" pageId="7" pageNumber="76">Incadendron esseri</emphasis>
</taxonomicName>
is given a preliminary status of Vulnerable (VU) under geographic range criteria B2 area of occupancy &lt;2000 km2 (B2a, known to exist at no more than 10 locations; B2b, continuing decline projected). Threats to this taxon in the Cordillera del
<normalizedToken originalValue="Cóndor">Condor</normalizedToken>
include mining for the underlying silica sand. Parts of its Peruvian range are protected within the Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
and Parque Nacional del Manu.
</paragraph>
</subSubSection>
<subSubSection lastPageId="8" lastPageNumber="77" pageId="7" pageNumber="76" type="additional collections">
<paragraph pageId="7" pageNumber="76">Additional collections.</paragraph>
<paragraph lastPageId="8" lastPageNumber="77" pageId="7" pageNumber="76">
<emphasis bold="true" pageId="7" pageNumber="76">ECUADOR. Zamora-Chinchipe</emphasis>
: Yantzaza
<normalizedToken originalValue="Cantón">Canton</normalizedToken>
. Cordillera del
<normalizedToken originalValue="Cóndor">Condor</normalizedToken>
region.
<normalizedToken originalValue="Río">Rio</normalizedToken>
Machinaza watershed, east of Los Encuentros, in and near a 0.25-ha forest inventory plot, tree #1477 in Aurelian Plot #6, La Zarza mining concession of Kinross Aurelian Corp., about 1.7 km southeast of and 500 m above Las
<normalizedToken originalValue="Peñas">Penas</normalizedToken>
camp,
<geoCoordinate degrees="03" direction="south" minutes="47" orientation="latitude" precision="15" seconds="50" value="-3.7972221">03°47'50&quot;S</geoCoordinate>
<geoCoordinate degrees="78" direction="west" minutes="29" orientation="longitude" precision="15" seconds="05" value="-78.48472">78°29'05&quot;W</geoCoordinate>
, 1840 m, 30 Jun 2009 (fl),
<emphasis italics="true" pageId="7" pageNumber="76">D. Neill &amp; C. Kajekai 16620</emphasis>
(MO, US); [same locality], tree #1362 in Aurelian Plot #6, 1840 m, 30 Jun 2009 (fl),
<emphasis italics="true" pageId="7" pageNumber="76">D. Neill &amp; C. Kajekai 16622</emphasis>
(MO, US); [same locality], tree #1477 in Aurelian Plot #6, 1840 m, 30 Jun 2009 (fl),
<emphasis italics="true" pageId="7" pageNumber="76">D. Neill &amp; C. Kajekai 16646</emphasis>
(MO, US).
<emphasis bold="true" pageId="7" pageNumber="76">PERU. Cusco</emphasis>
: Prov. Paucartambo,
<normalizedToken originalValue="Kosñipata">Kosnipata</normalizedToken>
, Trocha
<normalizedToken originalValue="Unión">Union</normalizedToken>
, km 13, Parcela VIII, subparcela 16,
<normalizedToken originalValue="árbol">arbol</normalizedToken>
706, 19L 0222887, UTM 8553630, 1835 m, 19 Aug 2003 (fl, fr),
<emphasis italics="true" pageId="7" pageNumber="76">W. Farfan et al. 1049</emphasis>
(CUZ, F, HUT, MO, USM, DAV; cited as
<emphasis italics="true" pageId="7" pageNumber="76">Garcia et al. 1049</emphasis>
in
<bibRefCitation DOI="https://doi.org/10.15381/rpb.v22i2.11351" author="Farfan-Rios, W" journalOrPublisher="Revista Peruana de Biologia" pageId="13" pageNumber="82" pagination="145 - 174" refId="B11" refString="Farfan-Rios, W, Garcia-Cabrera, K, Salinas, N, Silman, MR, 2015. Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando. Revista Peruana de Biologia 22: 145 - 174, DOI: https://doi.org/10.15381/rpb.v22i2.11351" title="Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando." url="https://doi.org/10.15381/rpb.v22i2.11351" volume="22" year="2015">Farfan-Rios et al. 2015</bibRefCitation>
); Trocha
<normalizedToken originalValue="Unión">Union</normalizedToken>
, km 10, parcela VI, subparcela 2,
<normalizedToken originalValue="árbol">arbol</normalizedToken>
74, 19L 0221737, UTM 8552556, 2295 m, 4 Sep 2003 (sterile),
<emphasis italics="true" pageId="7" pageNumber="76">W. Farfan et al. 1090</emphasis>
(CUZ, MO, USM, WFU); Trocha
<normalizedToken originalValue="Unión">Union</normalizedToken>
, km 11, parcela VII, subparcela 3,
<normalizedToken originalValue="árbol">arbol</normalizedToken>
105, 19L 0222622, UTM 8553538, 2000 m, 9 Sep 2003 (fl, fr),
<emphasis italics="true" pageId="7" pageNumber="76">W. Farfan et al. 1131</emphasis>
(CUZ, MO, USM, WFU); Prov. Paucartambo, Callanga, 19L 196221, UTM 8578219, 2245 m, 13 Sep 2007 (sterile),
<emphasis italics="true" pageId="7" pageNumber="76">W. Farfan et al. 3635</emphasis>
(USM, WFU); Callanga, 19L 196364, UTM 8579065, 2110 m, 14 Sep 2007 (sterile),
<emphasis italics="true" pageId="7" pageNumber="76">W. Farfan et al. 3696</emphasis>
(USM, WFU).
<emphasis bold="true" pageId="7" pageNumber="76">Pasco</emphasis>
: Oxapampa, Distrito Oxapampa, Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
,
<normalizedToken originalValue="cercanías">cercanias</normalizedToken>
del Refugio el Cedro,
<geoCoordinate degrees="10" direction="south" minutes="32" orientation="latitude" precision="925" value="-10.533334">10°32'S</geoCoordinate>
,
<geoCoordinate degrees="75" direction="west" minutes="21" orientation="longitude" precision="925" value="-75.35">75°21'W</geoCoordinate>
, 2240 m, 27 Nov 2002 (fl),
<emphasis italics="true" pageId="7" pageNumber="76">A. Monteagudo et al. 4458</emphasis>
(MO, US); Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
,
<normalizedToken originalValue="cercanías">cercanias</normalizedToken>
del Refugio el Cedro,
<geoCoordinate degrees="10" direction="south" minutes="32" orientation="latitude" precision="925" value="-10.533334">10°32'S</geoCoordinate>
,
<geoCoordinate degrees="75" direction="west" minutes="22" orientation="longitude" precision="925" value="-75.36667">75°22'W</geoCoordinate>
, 2200-2400 m, 6 Feb 2003 (fl, fr),
<emphasis italics="true" pageId="7" pageNumber="76">A. Monteagudo et al. 4484</emphasis>
(MO, US); Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
, camino del Refugio al Abra La Esperanza,
<geoCoordinate degrees="10" direction="south" minutes="31" orientation="latitude" precision="925" value="-10.516666">10°31'S</geoCoordinate>
,
<geoCoordinate degrees="75" direction="west" minutes="20" orientation="longitude" precision="925" value="-75.333336">75°20'W</geoCoordinate>
, 2400 m, 8 Mar 2003 (fr),
<emphasis italics="true" pageId="7" pageNumber="76">A. Monteagudo &amp; G. Ortiz 4605</emphasis>
(MO, US); Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
, Refugio Abra Esperanza y sus alrededores,
<geoCoordinate degrees="10" direction="south" minutes="31" orientation="latitude" precision="15" seconds="55" value="-10.531944">10°31'55&quot;S</geoCoordinate>
<geoCoordinate degrees="75" direction="west" minutes="20" orientation="longitude" precision="15" seconds="59" value="-75.349724">75°20'59&quot;W</geoCoordinate>
, 2786 m, 23 Apr 2009 (fr),
<emphasis italics="true" pageId="7" pageNumber="76">M. Cueva 592</emphasis>
(MO, US); Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
,
<normalizedToken originalValue="Estación">Estacion</normalizedToken>
<normalizedToken originalValue="Biológica">Biologica</normalizedToken>
San Alberto, Refugio El Cedro,
<geoCoordinate degrees="10" direction="south" minutes="32" orientation="latitude" precision="15" seconds="20" value="-10.538889">10°32'20&quot;S</geoCoordinate>
<geoCoordinate degrees="75" direction="west" minutes="20" orientation="longitude" precision="15" seconds="14" value="-75.33723">75°20'14&quot;W</geoCoordinate>
, 2731 m, 11 Feb 2012 (fr),
<emphasis italics="true" pageId="7" pageNumber="76">
R.
<normalizedToken originalValue="Vásquez">Vasquez</normalizedToken>
&amp; L. Valenzuela 37638
</emphasis>
(MO, US); Localidad
<normalizedToken originalValue="Grapanazú">Grapanazu</normalizedToken>
,
<geoCoordinate degrees="10" direction="south" minutes="29" orientation="latitude" precision="15" seconds="34" value="-10.492778">10°29'34&quot;S</geoCoordinate>
<geoCoordinate degrees="75" direction="west" minutes="23" orientation="longitude" precision="15" seconds="28" value="-75.391106">75°23'28&quot;W</geoCoordinate>
, 2288 m, 22 Nov 2012 (young fr),
<emphasis italics="true" pageId="7" pageNumber="76">
R.
<normalizedToken originalValue="Vásquez">Vasquez</normalizedToken>
38201
</emphasis>
(MO);
<pageBreakToken pageId="8" pageNumber="77" start="start">Distrito</pageBreakToken>
Huancabamba, zona de amortiguamiento del Parque Nacional Yanachaga
<normalizedToken originalValue="Chemillén">Chemillen</normalizedToken>
, al borde de las chacras y pastizales
<normalizedToken originalValue="cercanías">cercanias</normalizedToken>
de la casa del
<normalizedToken originalValue="Señor">Senor</normalizedToken>
Orlando Quispe,
<geoCoordinate degrees="10" direction="south" minutes="16" orientation="latitude" precision="15" seconds="38" value="-10.277222">10°16'38&quot;S</geoCoordinate>
<geoCoordinate degrees="75" direction="west" minutes="31" orientation="longitude" precision="15" seconds="06" value="-75.51833">75°31'06&quot;W</geoCoordinate>
, 1894 m, 24 Jul 2008 (young infl),
<emphasis italics="true" pageId="8" pageNumber="77">A. Monteagudo et al. 16929</emphasis>
(MO, US).
</paragraph>
</subSubSection>
<subSubSection lastPageId="9" lastPageNumber="78" pageId="8" pageNumber="77" type="discussion">
<paragraph pageId="8" pageNumber="77">Discussion.</paragraph>
<paragraph pageId="8" pageNumber="77">
Specimens of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
mostly were tentatively identified by collectors either as
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sapium" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Sapium" order="Malpighiales" pageId="8" pageNumber="77" phylum="Angiospermae" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Sapium</emphasis>
</taxonomicName>
Jacq., due to similarities in coriaceous leaves and glandular, spicate inflorescences (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="8" pageNumber="77">3A, D</figureCitation>
), or as
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Micrandra" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Micrandra" order="Malpighiales" pageId="8" pageNumber="77" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Micrandra</emphasis>
</taxonomicName>
Benth. (e.g.,
<bibRefCitation DOI="https://doi.org/10.15381/rpb.v22i2.11351" author="Farfan-Rios, W" journalOrPublisher="Revista Peruana de Biologia" pageId="13" pageNumber="82" pagination="145 - 174" refId="B11" refString="Farfan-Rios, W, Garcia-Cabrera, K, Salinas, N, Silman, MR, 2015. Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando. Revista Peruana de Biologia 22: 145 - 174, DOI: https://doi.org/10.15381/rpb.v22i2.11351" title="Lista anotada de arboles y afines en los bosques montanos del sureste peruano: la importancia de seguir recolectando." url="https://doi.org/10.15381/rpb.v22i2.11351" volume="22" year="2015">Farfan-Rios et al. 2015</bibRefCitation>
), due to their shared unusually large fruits and white latex.
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sapium" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Sapium" order="Malpighiales" pageId="8" pageNumber="77" phylum="Angiospermae" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Sapium</emphasis>
</taxonomicName>
notably differs in its bistaminate flowers and red-arillate seeds, and
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Micrandra" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Micrandra" order="Malpighiales" pageId="8" pageNumber="77" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Micrandra</emphasis>
</taxonomicName>
is florally very different as a member of subfamily
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="8" pageNumber="77" phylum="Magnoliophyta" rank="subFamily" subFamily="Crotonoideae">Crotonoideae</taxonomicName>
. Within the context of the generic key to South American Hippomaneae in
<bibRefCitation DOI="https://doi.org/10.1600/036364415X688781" author="Athie-Souza, SM" journalOrPublisher="Systematic Botany" pageId="13" pageNumber="82" pagination="527 - 533" refId="B1" refString="Athie-Souza, SM, Laurenio, de Melo A, Jose, da Silva M, dos, Santos Dias de Oliveira L, Ferreira, de Sales M, 2015. Gradyana (Euphorbiaceae): A new genus from northeastern Brazil. Systematic Botany 40: 527 - 533, DOI: https://doi.org/10.1600/036364415X688781" title="Gradyana (Euphorbiaceae): A new genus from northeastern Brazil." url="https://doi.org/10.1600/036364415X688781" volume="40" year="2015">
<normalizedToken originalValue="Athiê-Souza">Athie-Souza</normalizedToken>
et al. (2015)
</bibRefCitation>
,
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
would group with
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sebastiania" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Sebastiania" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Sebastiania</emphasis>
</taxonomicName>
Spreng. A comparison of select genera and distinguishing characters is given in Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Comparison of Incadendron with the morphologically most-similar neotropical genera. Based on primary observations with supplements from Kruijt (1996) and Esser (1995, 2001). The circumscription of Sebastiania is controversial leading to some uncertainty in the breadth of character states." httpUri="http://table.plazi.org/id/9FED5A1DB7703AE03E76D8FEAB57F42A" pageId="8" pageNumber="77" tableUuid="9FED5A1DB7703AE03E76D8FEAB57F42A">1</tableCitation>
. These genera are the most morphologically and geographically similar to
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
but are not necessarily its closest relatives, which are presently unclear.
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Senefelderopsis</emphasis>
</taxonomicName>
Steyerm., however, may have a closer relationship as suggested by similar fruit structure (see below), biogeographic ties between the Andean cordilleras and Guiana Shield (
<bibRefCitation author="Berry, PE" journalOrPublisher="Biologiske Skrifter" pageId="13" pageNumber="82" pagination="145 - 167" refId="B2" refString="Berry, PE, Riina, R, 2005. . Biologiske Skrifter 5: 145 - 167" volume="5" year="2005">Berry and Riina 2005</bibRefCitation>
), and isolated phylogenetic placement in the same diverse subclade H1 of Hippomaneae (
<bibRefCitation DOI="https://doi.org/10.3732/ajb.92.8.1397" author="Wurdack, KJ" journalOrPublisher="American Journal of Botany" pageId="14" pageNumber="83" pagination="1397 - 1420" refId="B25" refString="Wurdack, KJ, Hoffmann, P, Chase, MW, 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences. American Journal of Botany 92: 1397 - 1420, DOI: https://doi.org/10.3732/ajb.92.8.1397" title="Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences." url="https://doi.org/10.3732/ajb.92.8.1397" volume="92" year="2005">Wurdack et al. 2005</bibRefCitation>
).
</paragraph>
<paragraph pageId="8" pageNumber="77">
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
presents a unique combination of rare characters (discussed below) within Hippomaneae including sheathing stipules, conduplicate ptyxis, leaf margins entire and with unusual glands, and large, woody fruits. None of these characters appears phylogenetically very informative because they are autapomorphic or clearly homoplasious when considered in the context of molecular phylogenies (i.e.,
<bibRefCitation DOI="https://doi.org/10.3732/ajb.92.8.1397" author="Wurdack, KJ" journalOrPublisher="American Journal of Botany" pageId="14" pageNumber="83" pagination="1397 - 1420" refId="B25" refString="Wurdack, KJ, Hoffmann, P, Chase, MW, 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences. American Journal of Botany 92: 1397 - 1420, DOI: https://doi.org/10.3732/ajb.92.8.1397" title="Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences." url="https://doi.org/10.3732/ajb.92.8.1397" volume="92" year="2005">Wurdack et al. 2005</bibRefCitation>
, Wurdack, unpublished). Thus, while the rare character combination serves well to distinguish
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
and indicates a degree of morphological disparity deserving of generic recognition, it does not inform on relationships nor provide much insight into how such characters evolved. Major floral features that are variable in Hippomaneae have broadly distributed, likely plesiomorphic, states in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
, including terminal and unbranched inflorescences, single pistillate flower per bisexual inflorescence, staminate cymules that are multiflowered and glandular, and 3-merous pedicellate staminate flowers with free stamens.
</paragraph>
<paragraph lastPageId="9" lastPageNumber="78" pageId="8" pageNumber="77">
Pollen morphology and ultrastructure are remarkably diverse across
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" family="Euphorbiaceae" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="8" pageNumber="77" rank="family">Euphorbiaceae</taxonomicName>
, but are less variable in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" family="Euphorbiaceae" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="8" pageNumber="77" rank="subFamily" subFamily="Euphorbioideae">Euphorbioideae</taxonomicName>
, and Hippomaneae in particular (
<bibRefCitation DOI="https://doi.org/10.1111/j.1438-8677.1962.tb00010.x" author="Punt, W" journalOrPublisher="Wentia" pageId="14" pageNumber="83" pagination="1 - 116" refId="B19" refString="Punt, W, 1962. Pollen morphology of the Euphorbiaceae with special reference to taxonomy. Wentia 7: 1 - 116, DOI: https://doi.org/10.1111/j.1438-8677.1962.tb00010.x" title="Pollen morphology of the Euphorbiaceae with special reference to taxonomy." url="https://doi.org/10.1111/j.1438-8677.1962.tb00010.x" volume="7" year="1962">Punt 1962</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.11110/kjpt.2013.43.1.1" author="Park, K-R" journalOrPublisher="Korean Journal of Plant Taxonomy" pageId="14" pageNumber="83" pagination="1 - 11" refId="B18" refString="Park, K-R, Lee, J-H, 2013. Pollen morphology of subfamily Euphorbioideae (Euphorbiaceae). Korean Journal of Plant Taxonomy 43: 1 - 11, DOI: https://doi.org/10.11110/kjpt.2013.43.1.1" title="Pollen morphology of subfamily Euphorbioideae (Euphorbiaceae)." url="https://doi.org/10.11110/kjpt.2013.43.1.1" volume="43" year="2013">Park and Lee 2013</bibRefCitation>
). The pollen of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
resembles that of other hippomanoids in being tricolporate with a perforate exine (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="8" pageNumber="77">4C-D</figureCitation>
). The seed structure of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
is also similar to that of other Hippomaneae (
<bibRefCitation DOI="https://doi.org/10.2307/2421457" author="Martin, AC" journalOrPublisher="The American Midland Naturalist" pageId="14" pageNumber="83" pagination="513 - 660" refId="B16" refString="Martin, AC, 1946. The comparative internal morphology of seeds. The American Midland Naturalist 36: 513 - 660, DOI: https://doi.org/10.2307/2421457" title="The comparative internal morphology of seeds." url="https://doi.org/10.2307/2421457" volume="36" year="1946">Martin 1946</bibRefCitation>
,
<bibRefCitation DOI="https://doi.org/10.1007/s10265-002-0047-5" author="Tokuoka, T" journalOrPublisher="Journal of Plant Research" pageId="14" pageNumber="83" pagination="361 - 374" refId="B22" refString="Tokuoka, T, Tobe, H, 2002. Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications. Journal of Plant Research 115: 361 - 374, DOI: https://doi.org/10.1007/s10265-002-0047-5" title="Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications." url="https://doi.org/10.1007/s10265-002-0047-5" volume="115" year="2002">Tokuoka and Tobe 2002</bibRefCitation>
). Its seed coat (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="8" pageNumber="77">3L</figureCitation>
) with a thin testa of collapsed cells and palisade-like mechanical exotegmen of elongate, pitted, thick-walled cells resembles that of exotegmic genera across the entire family. Depending on exact location, the pallisadal cells vary in length (3-4
<normalizedToken originalValue="×">x</normalizedToken>
shorter at apex versus bottom and sides) and orientation (vertical, inclined, or curved). Embryos generally deserve more careful description and study given their diversity in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" family="Euphorbiaceae" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="8" pageNumber="77" rank="family">Euphorbiaceae</taxonomicName>
, although most are variants of the spathulate fully developed type (
<bibRefCitation DOI="https://doi.org/10.2307/2421457" author="Martin, AC" journalOrPublisher="The American Midland Naturalist" pageId="14" pageNumber="83" pagination="513 - 660" refId="B16" refString="Martin, AC, 1946. The comparative internal morphology of seeds. The American Midland Naturalist 36: 513 - 660, DOI: https://doi.org/10.2307/2421457" title="The comparative internal morphology of seeds." url="https://doi.org/10.2307/2421457" volume="36" year="1946">Martin 1946</bibRefCitation>
). The embryo of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="8" pageNumber="77" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="8" pageNumber="77">Incadendron</emphasis>
</taxonomicName>
<pageBreakToken pageId="9" pageNumber="78" start="start">agrees</pageBreakToken>
with this generalization (see Description for details, based on one intact seed) and in particular resembles that of other Hippomaneae including
<taxonomicName authorityName="Steyerm" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis croizatii" order="Malpighiales" pageId="9" pageNumber="78" phylum="Tracheophyta" rank="species" species="croizatii">
<emphasis italics="true" pageId="9" pageNumber="78">Senefelderopsis croizatii</emphasis>
</taxonomicName>
Steyerm. (examined here,
<emphasis italics="true" pageId="9" pageNumber="78">Maas et al. 5828</emphasis>
, US), which has a slightly longer hypocotyl-radicle axis at 3
<normalizedToken originalValue="×">x</normalizedToken>
1 mm (versus 2
<normalizedToken originalValue="×">x</normalizedToken>
1 mm) and thicker cotyledons at 0.5 mm (versus 0.1 mm).
</paragraph>
</subSubSection>
<subSubSection lastPageId="12" lastPageNumber="81" pageId="9" pageNumber="78" type="unusual morphological features of incadendron">
<paragraph pageId="9" pageNumber="78">
Unusual morphological features of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="9" pageNumber="78" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="78">Incadendron</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph lastPageId="10" lastPageNumber="79" pageId="9" pageNumber="78">
The entire hippomanoid clade (sensu
<bibRefCitation DOI="https://doi.org/10.3732/ajb.92.8.1397" author="Wurdack, KJ" journalOrPublisher="American Journal of Botany" pageId="14" pageNumber="83" pagination="1397 - 1420" refId="B25" refString="Wurdack, KJ, Hoffmann, P, Chase, MW, 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences. American Journal of Botany 92: 1397 - 1420, DOI: https://doi.org/10.3732/ajb.92.8.1397" title="Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euphorbiaceae sensu stricto) using plastid rbcL and trnL-F DNA sequences." url="https://doi.org/10.3732/ajb.92.8.1397" volume="92" year="2005">Wurdack et al. 2005</bibRefCitation>
) is considered here for the purposes of broad discussion rather than restricted to paraphyletic Hippomaneae. This broad grouping is reflected in the nomenclaturally problematic (see
<bibRefCitation DOI="https://doi.org/10.1590/S2175-78602012000100013" author="Esser, H-J" journalOrPublisher="Rodriguesia" pageId="13" pageNumber="82" pagination="209 - 225" refId="B8" refString="Esser, H-J, 2012. The tribe Hippomaneae (Euphorbiaceae) in Brazil. Rodriguesia 63: 209 - 225, DOI: https://doi.org/10.1590/S2175-78602012000100013" title="The tribe Hippomaneae (Euphorbiaceae) in Brazil." url="https://doi.org/10.1590/S2175-78602012000100013" volume="63" year="2012">Esser 2012</bibRefCitation>
) Hippomaneae s.l. of
<bibRefCitation DOI="https://doi.org/10.1007/978-3-642-39417-1_10" author="Webster, GL" editor="Kubitzki, K" journalOrPublisher="Flowering plants. Eudicots. Malpighiales [Vol. 11]. Springer-Verlag, Berlin Heidelberg, Germany" pageId="14" pageNumber="83" pagination="51 - 216" refId="B24" refString="Webster, GL, 2014. Euphorbiaceae. In: Kubitzki, K, Ed., The Families and Genera of Vascular Plants. Flowering plants. Eudicots. Malpighiales [Vol. 11]. Springer-Verlag, Berlin Heidelberg, Germany: 51 - 216, DOI: https://doi.org/10.1007/978-3-642-39417-1_10" title="Euphorbiaceae." url="https://doi.org/10.1007/978-3-642-39417-1_10" volumeTitle="The Families and Genera of Vascular Plants." year="2014">Webster (2014)</bibRefCitation>
. Stipules in the hippomanoids are typically small, scale-like (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Hippomaneae shoot tips showing stipule and ptyxis variation. A Homalanthus nutans (G. Forst.) Guill. with large, deciduous, sheathing stipules protecting shoot tip B Transverse view of H. nutans shoot tip, showing nested series of three developing leaves with scroll-like lamina (ptyxis involute), each surrounded by a pair of sheathing stipules C Transverse view of Excoecaria cochinchinensis Lour. young leaf showing slightly inrolled lamina (ptyxis conduplicate-involute) D E. cochinchinensis shoot tip with small persistent stipules. (Abbreviation: s = stipule. Source: Freehand sections of fresh tissues grown in Department of Botany greenhouses)." figureDoi="10.3897/phytokeys.85.14757.figure5" httpUri="https://binary.pensoft.net/fig/153295" pageId="9" pageNumber="78">5D</figureCitation>
) or absent. Conspicuous, sheathing stipules are relatively rare and in addition to
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="9" pageNumber="78" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="78">Incadendron</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Illustration of Incadendron esseri. A Habit B Shoot tip C Leaf base (adaxial) D Leaf base and marginal glands (abaxial). E Staminate subinflorescence F Staminate cymule (distal view) G Staminate cymule (proximal view, without lateral buds) H Staminate flower I Pistillate flower J Fruit K Mericarp valve L Columella M Seed (ventral face). (Source: A-G Calatayud et al. 4711, MO; H-I Monteagudo et al. 4458, US; J Vasquez &amp; Valenzuela 37638, MO; K-M Monteagudo &amp; Ortiz 4605, US)." figureDoi="10.3897/phytokeys.85.14757.figure1" httpUri="https://binary.pensoft.net/fig/153291" pageId="9" pageNumber="78">1B</figureCitation>
) also characterize
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Conosapium" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Conosapium" order="Malpighiales" pageId="9" pageNumber="78" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="78">Conosapium</emphasis>
</taxonomicName>
<normalizedToken originalValue="Müll">Muell</normalizedToken>
. Arg.,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Homalanthus" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Homalanthus" order="Malpighiales" pageId="9" pageNumber="78" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="9" pageNumber="78">Homalanthus</emphasis>
</taxonomicName>
A. Juss. (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Hippomaneae shoot tips showing stipule and ptyxis variation. A Homalanthus nutans (G. Forst.) Guill. with large, deciduous, sheathing stipules protecting shoot tip B Transverse view of H. nutans shoot tip, showing nested series of three developing leaves with scroll-like lamina (ptyxis involute), each surrounded by a pair of sheathing stipules C Transverse view of Excoecaria cochinchinensis Lour. young leaf showing slightly inrolled lamina (ptyxis conduplicate-involute) D E. cochinchinensis shoot tip with small persistent stipules. (Abbreviation: s = stipule. Source: Freehand sections of fresh tissues grown in Department of Botany greenhouses)." figureDoi="10.3897/phytokeys.85.14757.figure5" httpUri="https://binary.pensoft.net/fig/153295" pageId="9" pageNumber="78">5A</figureCitation>
),
<taxonomicName class="Equisetopsida" family="Euphorbiaceae" genus="Hura" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Hura" order="Malpighiales" pageId="10" pageNumber="79" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">
<pageBreakToken pageId="10" pageNumber="79" start="start">Hura</pageBreakToken>
</emphasis>
</taxonomicName>
L.,
<taxonomicName authorityName="H.-J.Esser" authorityYear="1998" class="Magnoliopsida" family="Euphorbiaceae" genus="Neoshirakia" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Neoshirakia" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Neoshirakia</emphasis>
</taxonomicName>
Esser, and
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Pachystroma" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Pachystroma" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Pachystroma</emphasis>
</taxonomicName>
<normalizedToken originalValue="Müll">Muell</normalizedToken>
. Arg. While functionally these large stipules are similar in conferring additional protection to the shoot apex, they differ in morphological details and likely are convergent size increases. The stipules in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Incadendron</emphasis>
</taxonomicName>
are large, sheath the terminal bud, unusually stout in keeping with the coriaceous leaves, deciduous, and centrally attached such that an elliptic scar remains after abscission (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="10" pageNumber="79">3E</figureCitation>
). The buds are internally mucilaginous. Serial sections of a single bud show a distinct palisade-like cell layer on the inner surfaces of the stipules that is the likely source of this secretion (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="10" pageNumber="79">4H-I</figureCitation>
). This layer appears to differentiate late in development as it is present in the outer stipules but not on the enclosed next younger pair of the bud (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="10" pageNumber="79">4H</figureCitation>
). The inflorescence bracts resemble a smaller version of the stipules (rarely with a few subtending glands of the type found with the staminate cymules) and are also deciduous, leaving large scars (Figs
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="10" pageNumber="79">3F</figureCitation>
,
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="10" pageNumber="79">4B</figureCitation>
). Conspicuous sheathing stipules in other taxa are mostly thinner except in
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Pachystroma" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Pachystroma" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Pachystroma</emphasis>
</taxonomicName>
, and may be centrally attached, leaving elliptic scars (
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Homalanthus" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Homalanthus" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Homalanthus</emphasis>
</taxonomicName>
,
<taxonomicName class="Equisetopsida" family="Euphorbiaceae" genus="Hura" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Hura" order="Malpighiales" pageId="10" pageNumber="79" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Hura</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="H.-J.Esser" authorityYear="1998" class="Magnoliopsida" family="Euphorbiaceae" genus="Neoshirakia" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Neoshirakia" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Neoshirakia</emphasis>
</taxonomicName>
) or broadly attached along their entire base, leaving semi-circumferential scars (
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Conosapium" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Conosapium" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Conosapium</emphasis>
</taxonomicName>
,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Pachystroma" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Pachystroma" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Pachystroma</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph lastPageId="11" lastPageNumber="80" pageId="10" pageNumber="79">
Leaf folding (ptyxis) is variable but poorly studied for
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="family">Euphorbiaceae</taxonomicName>
in general (
<bibRefCitation author="Cullen, J" journalOrPublisher="Notes from the Royal Botanic Garden Edinburgh" pageId="13" pageNumber="82" pagination="161 - 214" refId="B4" refString="Cullen, J, 1978. A preliminary survey of ptyxis (vernation) in angiosperms. Notes from the Royal Botanic Garden Edinburgh 37: 161 - 214" title="A preliminary survey of ptyxis (vernation) in angiosperms." volume="37" year="1978">Cullen 1978</bibRefCitation>
). Based on recent surveys (K. Wurdack, personal observations)
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="10" pageNumber="79" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="10" pageNumber="79">Incadendron</emphasis>
</taxonomicName>
appears unique in the hippomanoids in having conduplicate ptyxis (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="10" pageNumber="79">4G</figureCitation>
). In bud and early expansion after stipule abscission, the halves of the blade are folded tightly together along their adaxial surface and the leaf in transverse section shows no curling at the edges. After the blade halves spread open, the margins recurve before hardening and finally at maturity the halves assume a flattened aspect with a slightly revolute margin (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="10" pageNumber="79">3A, C</figureCitation>
). The terminal buds are flattened (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="10" pageNumber="79">3E</figureCitation>
,
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="10" pageNumber="79">4G</figureCitation>
), which likely reflects the conduplicate nature of the enclosed developing leaf blades, although this point needs further study with fresh material. Other hippomanoids have varying
<pageBreakToken pageId="11" pageNumber="80" start="start">degrees</pageBreakToken>
of developing leaf blade curvature including tightly rolled scrolls, rolled edges, or gently curved loops that span involute (e.g.,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Homalanthus" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Homalanthus" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Homalanthus</emphasis>
</taxonomicName>
, Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Hippomaneae shoot tips showing stipule and ptyxis variation. A Homalanthus nutans (G. Forst.) Guill. with large, deciduous, sheathing stipules protecting shoot tip B Transverse view of H. nutans shoot tip, showing nested series of three developing leaves with scroll-like lamina (ptyxis involute), each surrounded by a pair of sheathing stipules C Transverse view of Excoecaria cochinchinensis Lour. young leaf showing slightly inrolled lamina (ptyxis conduplicate-involute) D E. cochinchinensis shoot tip with small persistent stipules. (Abbreviation: s = stipule. Source: Freehand sections of fresh tissues grown in Department of Botany greenhouses)." figureDoi="10.3897/phytokeys.85.14757.figure5" httpUri="https://binary.pensoft.net/fig/153295" pageId="11" pageNumber="80">5B</figureCitation>
) to supervolute-curved ptyxis (e.g.,
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefelderopsis</emphasis>
</taxonomicName>
).
<bibRefCitation author="Cullen, J" journalOrPublisher="Notes from the Royal Botanic Garden Edinburgh" pageId="13" pageNumber="82" pagination="161 - 214" refId="B4" refString="Cullen, J, 1978. A preliminary survey of ptyxis (vernation) in angiosperms. Notes from the Royal Botanic Garden Edinburgh 37: 161 - 214" title="A preliminary survey of ptyxis (vernation) in angiosperms." volume="37" year="1978">Cullen (1978)</bibRefCitation>
indicated the hippomanoid
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Excoecaria" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Excoecaria cochinchinensis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="cochinchinensis">
<emphasis italics="true" pageId="11" pageNumber="80">Excoecaria cochinchinensis</emphasis>
</taxonomicName>
Lour. (as
<taxonomicName lsidName="E. bicolor" pageId="11" pageNumber="80" rank="species" species="bicolor">
<emphasis italics="true" pageId="11" pageNumber="80">E. bicolor</emphasis>
</taxonomicName>
[Hassk.] Zoll. ex Hassk.) was conduplicate. Our observations on living (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Hippomaneae shoot tips showing stipule and ptyxis variation. A Homalanthus nutans (G. Forst.) Guill. with large, deciduous, sheathing stipules protecting shoot tip B Transverse view of H. nutans shoot tip, showing nested series of three developing leaves with scroll-like lamina (ptyxis involute), each surrounded by a pair of sheathing stipules C Transverse view of Excoecaria cochinchinensis Lour. young leaf showing slightly inrolled lamina (ptyxis conduplicate-involute) D E. cochinchinensis shoot tip with small persistent stipules. (Abbreviation: s = stipule. Source: Freehand sections of fresh tissues grown in Department of Botany greenhouses)." figureDoi="10.3897/phytokeys.85.14757.figure5" httpUri="https://binary.pensoft.net/fig/153295" pageId="11" pageNumber="80">5C</figureCitation>
) and herbarium material indicate its glandular-toothed margin is inwardly curled and the ptyxis is more accurately described as a conduplicate-involute intermediate. The hippomanoids contain a wide diversity of petiolar and leaf gland form and position.
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
has no acropetiolar or embedded laminar glands but it has glands along the leaf margin, which are best developed (i.e., consistent in presence and largest in size) near the leaf base. Curling of the leaf edge with age shields these basal glands (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="11" pageNumber="80">3C</figureCitation>
), although the scattered more distal glands remain exposed. The marginal glands (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="11" pageNumber="80">4E</figureCitation>
) are not associated with teeth or setae and are similar in morphology to those found on
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Gymnanthes" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Gymnanthes schottiana" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="schottiana">
<emphasis italics="true" pageId="11" pageNumber="80">Gymnanthes schottiana</emphasis>
</taxonomicName>
<normalizedToken originalValue="Müll">Muell</normalizedToken>
. Arg. Hippomanoid leaf margins typically have regular teeth or marginal setae, and are more rarely similar to
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
or entire without any associated glands. The abaxial leaf surface of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
is finely striate, but the stomata are not shielded or sunken (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Micromorphological and anatomical features of Incadendron. A Staminate flower with one anther removed, showing short filaments and basally connate sepals B Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts C Pollen; mesocolpium-centered equatorial view of whole grain D Pollen; slightly oblique polar view of whole grains E Abaxial laminar surface with marginal gland F Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata G Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis H Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image) I Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule. A-G imaged with a Zeiss EVO MA 15 SEM at 10 - 12 kV after sputter coating with 25 nm of Au / Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers. H-I from paraffin-embedded, rehydrated herbarium specimens; 7 μm sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source: A, C-D Monteagudo et al. 4458, US; B Neill &amp; Kajekai 16622, US; E-I Neill &amp; Kajekai 16646, US)." figureDoi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="11" pageNumber="80">4F</figureCitation>
). Some high elevation hippomanoids have micro-papillose surfaces and concealed stomata (e.g.,
<taxonomicName authorityName="H.-J.Esser" authorityYear="1993" class="Magnoliopsida" family="Euphorbiaceae" genus="Dendrothrix" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Dendrothrix" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Dendrothrix</emphasis>
</taxonomicName>
Esser,
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefelderopsis</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph pageId="11" pageNumber="80">
Inflorescences in the hippomanoids are axillary and/or terminal. In
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
the inflorescence is terminal but appears distinctly leaf-opposed due to near simultaneous development of both the inflorescence and adjacent leaf, coupled with the start of renewal shoot growth from the axillary bud before flowering is finished (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="11" pageNumber="80">3E</figureCitation>
). Axial growth and orientation in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
branches is not deflected by inflorescence development and gives little hint of being sympodial by substitution. Specimens of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
show twinned branching (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Illustration of Incadendron esseri. A Habit B Shoot tip C Leaf base (adaxial) D Leaf base and marginal glands (abaxial). E Staminate subinflorescence F Staminate cymule (distal view) G Staminate cymule (proximal view, without lateral buds) H Staminate flower I Pistillate flower J Fruit K Mericarp valve L Columella M Seed (ventral face). (Source: A-G Calatayud et al. 4711, MO; H-I Monteagudo et al. 4458, US; J Vasquez &amp; Valenzuela 37638, MO; K-M Monteagudo &amp; Ortiz 4605, US)." figureDoi="10.3897/phytokeys.85.14757.figure1" httpUri="https://binary.pensoft.net/fig/153291" pageId="11" pageNumber="80">1A</figureCitation>
,
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="11" pageNumber="80">3A</figureCitation>
), due to the occasional growth of additional leaf-axil accessory buds close to the growing branch tips. Most accessory buds, even in older parts of the shoot system, remain as barely visible meristems.
</paragraph>
<paragraph pageId="11" pageNumber="80">
Large fruits, ≥2 cm in diameter are relatively rare among the hippomanoids (i.e.,
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron esseri" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="esseri">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron esseri</emphasis>
</taxonomicName>
,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Duvigneaudia" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Duvigneaudia inopinata" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="inopinata">
<emphasis italics="true" pageId="11" pageNumber="80">Duvigneaudia inopinata</emphasis>
</taxonomicName>
[Prain] J.
<normalizedToken originalValue="Léonard">Leonard</normalizedToken>
,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Hippomane" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Hippomane mancinella" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="mancinella">
<emphasis italics="true" pageId="11" pageNumber="80">Hippomane mancinella</emphasis>
</taxonomicName>
L.,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Pachystroma" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Pachystroma longifolium" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="longifolium">
<emphasis italics="true" pageId="11" pageNumber="80">Pachystroma longifolium</emphasis>
</taxonomicName>
[Nees] I.M. Johnst.,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sebastiania" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Sebastiania obtusifolia" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="obtusifolia">
<emphasis italics="true" pageId="11" pageNumber="80">Sebastiania obtusifolia</emphasis>
</taxonomicName>
Pax &amp; K. Hoffm., and a few species each of
<taxonomicName authorityName="Baillon" authorityYear="1858" class="Magnoliopsida" family="Euphorbiaceae" genus="Algernonia" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Algernonia" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Algernonia</emphasis>
</taxonomicName>
Baill.,
<taxonomicName authorityName="Linnaeus" authorityYear="1759" class="Magnoliopsida" family="Euphorbiaceae" genus="Excoecaria" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Excoecaria" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Excoecaria</emphasis>
</taxonomicName>
L.,
<taxonomicName class="Equisetopsida" family="Euphorbiaceae" genus="Hura" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Hura" order="Malpighiales" pageId="11" pageNumber="80" phylum="Magnoliophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Hura</emphasis>
</taxonomicName>
L.,
<taxonomicName authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Euphorbiaceae" genus="Mabea" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Mabea" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Mabea</emphasis>
</taxonomicName>
Aubl.,
<taxonomicName authorityName="Freire Allemao" authorityYear="1849" class="Magnoliopsida" family="Euphorbiaceae" genus="Ophthalmoblapton" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Ophthalmoblapton" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Ophthalmoblapton</emphasis>
</taxonomicName>
<normalizedToken originalValue="Allemão">Allemao</normalizedToken>
,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sapium" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Sapium" order="Malpighiales" pageId="11" pageNumber="80" phylum="Angiospermae" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Sapium</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="C.F.P.Martius" authorityYear="1841" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefeldera" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefeldera" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefeldera</emphasis>
</taxonomicName>
Mart.,
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefelderopsis</emphasis>
</taxonomicName>
, and
<taxonomicName authorityName="H.-J.Esser" authorityYear="1999" class="Magnoliopsida" family="Euphorbiaceae" genus="Shirakiopsis" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Shirakiopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Shirakiopsis</emphasis>
</taxonomicName>
Esser), which otherwise typically have fruits less than half that size. Among the likely correlates of increased fruit size are larger seeds (see size variation in
<bibRefCitation DOI="https://doi.org/10.1007/s10265-002-0047-5" author="Tokuoka, T" journalOrPublisher="Journal of Plant Research" pageId="14" pageNumber="83" pagination="361 - 374" refId="B22" refString="Tokuoka, T, Tobe, H, 2002. Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications. Journal of Plant Research 115: 361 - 374, DOI: https://doi.org/10.1007/s10265-002-0047-5" title="Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications." url="https://doi.org/10.1007/s10265-002-0047-5" volume="115" year="2002">Tokuoka and Tobe 2002</bibRefCitation>
), thickened pericarps, indehiscence, and being borne singly (rarely two). Especially noteworthy is the combination of a dehiscent fruit with a thick pericarp (≥3 mm), which is rare among the big-fruited species. These generalizations do have exceptions such as thin walls in inflated fruits (i.e.,
<taxonomicName authorityName="Pax" authorityYear="1912" baseAuthorityName="Pax" class="Magnoliopsida" family="Euphorbiaceae" genus="Excoecaria" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Excoecaria bussei" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="species" species="bussei">
<emphasis italics="true" pageId="11" pageNumber="80">Excoecaria bussei</emphasis>
</taxonomicName>
[Pax] Pax), or multiple fruits in large-fruited taxa with robust or compound inflorescences (e.g.,
<emphasis italics="true" pageId="11" pageNumber="80">
<taxonomicName authorityName="Aublet" authorityYear="1775" class="Magnoliopsida" family="Euphorbiaceae" genus="Mabea" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Mabea" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">Mabea</taxonomicName>
<taxonomicName authorityName="C.F.P.Martius" authorityYear="1841" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefeldera" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefeldera" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">Senefeldera</taxonomicName>
</emphasis>
,
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefelderopsis</emphasis>
</taxonomicName>
).
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
exhibits some of these large-fruit trends including seed size, pericarp thickness, and being borne singly.
</paragraph>
<paragraph lastPageId="12" lastPageNumber="81" pageId="11" pageNumber="80">
The fruits of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Incadendron</emphasis>
</taxonomicName>
most closely resemble those of
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="11" pageNumber="80" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="11" pageNumber="80">Senefelderopsis</emphasis>
</taxonomicName>
. In addition to large size, they share many structural features including a thick pericarp with an ex
<pageBreakToken pageId="12" pageNumber="81" start="start">ceptionally</pageBreakToken>
thick woody mesocarp (≥3 mm), sharp woody apex, well-developed septa, mericarp valves connected with a basal triangle, and a thin funicle. These features individually (or in various combinations) occur in other genera, but the thick mesocarp is exceptional among the large-fruited, dehiscent species. The mesocarp of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
has a prominent dorsal-suture lip, has non-vascularized raised ridges that follow underneath major exocarp venation, and is stratified with a darker layer of different structure lining the locule (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81">3M</figureCitation>
); these features are to varying degrees shared with
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81">3N</figureCitation>
). In both
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
the robust septa dividing the locules are lignified and nearly complete except for a small apical gap (Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81">3J-K, M-N</figureCitation>
) to accommodate the funicle and its attachment to the placenta. Many other hippomanoids differ in having thin, poorly lignified septa that become destroyed during dehiscence, and/or septa that are incomplete leaving a large gap descriptively called a &quot;C-shaped cut&quot; (
<bibRefCitation DOI="https://doi.org/10.1600/036364415X688781" author="Athie-Souza, SM" journalOrPublisher="Systematic Botany" pageId="13" pageNumber="82" pagination="527 - 533" refId="B1" refString="Athie-Souza, SM, Laurenio, de Melo A, Jose, da Silva M, dos, Santos Dias de Oliveira L, Ferreira, de Sales M, 2015. Gradyana (Euphorbiaceae): A new genus from northeastern Brazil. Systematic Botany 40: 527 - 533, DOI: https://doi.org/10.1600/036364415X688781" title="Gradyana (Euphorbiaceae): A new genus from northeastern Brazil." url="https://doi.org/10.1600/036364415X688781" volume="40" year="2015">
<normalizedToken originalValue="Athiê-Souza">Athie-Souza</normalizedToken>
et al. 2015
</bibRefCitation>
). Fruit vascular variation, especially septal strand number, has been considered potentially informative at the generic level in Hippomaneae (
<bibRefCitation DOI="https://doi.org/10.2307/2419431" author="Esser, H-J" journalOrPublisher="Systematic Botany" pageId="13" pageNumber="82" pagination="617 - 628" refId="B9" refString="Esser, H-J, van Welzen, P, Djarwaningsih, T, 1997. A phylogenetic classification of the Malesian Hippomaneae (Euphorbiaceae). Systematic Botany 22: 617 - 628, DOI: https://doi.org/10.2307/2419431" title="A phylogenetic classification of the Malesian Hippomaneae (Euphorbiaceae)." url="https://doi.org/10.2307/2419431" volume="22" year="1997">Esser et al. 1997</bibRefCitation>
,
<bibRefCitation author="Esser, H-J" editor="Radcliffe-Smith, A" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="13" pageNumber="82" pagination="352 - 98" refId="B7" refString="Esser, H-J, 2001. Tribe 49. Hippomaneae. In: Radcliffe-Smith, A, Ed., Genera Euphorbiacearum. Royal Botanic Gardens, Kew, UK: 352 - 98" title="Tribe 49. Hippomaneae." volumeTitle="Genera Euphorbiacearum." year="2001">Esser 2001</bibRefCitation>
), but can be difficult to observe and its functional significance is not well understood. The septal vascular strand numbers are very similar between
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
(single or bifurcating) and
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(single or rarely bifurcating). The seemingly delicate, suspended nature of the relatively large seeds in the locules of both genera is unusual. Their thin, nearly erect funicles differ in length, which allow the seeds of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
to be further displaced downward in the locule from the point of attachment to the columella as compared with
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81">3I-K</figureCitation>
). Many other hippomanoid taxa have better support for their seeds either through greater filling of the locule cavity by the seed that leaves little free space around it, and/or through more robust attachment to the columella via a short and thickened funicle or fusion along the raphe. The fruit and locule shape differ slightly between the genera in being subglobose in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
versus elongate and more trigonous in
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Morphology of Incadendron (A-J, L-M) and Senefelderopsis (K, N). A Habit, with paired branching and staminate inflorescences; note latex at damaged nodes B Leaf base (adaxial) with basal lobes C Leaf base (abaxial) with curled glandular margin D Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule E Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot F Summit of peduncle showing bract scars G Nearly mature green fruit H Mature brown fruit I Seed with funicle; holes in I &amp; J come from insect predation J Mericarp valve with outline of seed position K Mericarp valve with a seed; funicle obscures gap L Seed coat, transverse view (SEM) M Pericarp profile and top half of valve (exocarp removed) N Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of M-N relative to J-K shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source: Incadendron, A-E, G-H Farfan et al. 1049, 1131; F Farfan et al. 706, MO; I-J, M Monteagudo &amp; Ortiz 4605, US; L Monteagudo et al. 4484, US. Senefelderopsis croizatii, K, N Radosavljevic 296, US)." figureDoi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81">3J-K, M-N</figureCitation>
). In addition, the exocarps (not examined anatomically and topographically defined here as the more or less removable outer layer) differ in gross structure with prominent vasculature at the interface between the loosely attached exocarp and woody mesocarp in
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
, likely supporting greater fleshiness when fresh. In
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
there is little exocarp vascularization and tight adherence via a stiff, porous layer of apparently sclerified cells that is best developed in
<taxonomicName lsidName="S. chiribiquetensis" pageId="12" pageNumber="81" rank="species" species="chiribiquetensis">
<emphasis italics="true" pageId="12" pageNumber="81">S. chiribiquetensis</emphasis>
</taxonomicName>
(R.E. Schult. &amp; Croizat) Steyerm. Although there are considerable morphological differences between
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
and
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(see Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Comparison of Incadendron with the morphologically most-similar neotropical genera. Based on primary observations with supplements from Kruijt (1996) and Esser (1995, 2001). The circumscription of Sebastiania is controversial leading to some uncertainty in the breadth of character states." httpUri="http://table.plazi.org/id/9FED5A1DB7703AE03E76D8FEAB57F42A" pageId="12" pageNumber="81" tableUuid="9FED5A1DB7703AE03E76D8FEAB57F42A">1</tableCitation>
), the striking fruit similarities may have special significance for indicating a closer relationship between the genera.
</paragraph>
<caption ID-Zenodo-Dep="995265" doi="10.3897/phytokeys.85.14757.figure1" httpUri="https://binary.pensoft.net/fig/153291" pageId="12" pageNumber="81" start="Figure 1" startId="F1">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Figure 1.</emphasis>
Illustration of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron esseri" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="esseri">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron esseri</emphasis>
</taxonomicName>
.
<emphasis bold="true" pageId="12" pageNumber="81">A</emphasis>
Habit
<emphasis bold="true" pageId="12" pageNumber="81">B</emphasis>
Shoot tip
<emphasis bold="true" pageId="12" pageNumber="81">C</emphasis>
Leaf base (adaxial)
<emphasis bold="true" pageId="12" pageNumber="81">D</emphasis>
Leaf base and marginal glands (abaxial).
<emphasis bold="true" pageId="12" pageNumber="81">E</emphasis>
Staminate subinflorescence
<emphasis bold="true" pageId="12" pageNumber="81">F</emphasis>
Staminate cymule (distal view)
<emphasis bold="true" pageId="12" pageNumber="81">G</emphasis>
Staminate cymule (proximal view, without lateral buds)
<emphasis bold="true" pageId="12" pageNumber="81">H</emphasis>
Staminate flower
<emphasis bold="true" pageId="12" pageNumber="81">I</emphasis>
Pistillate flower
<emphasis bold="true" pageId="12" pageNumber="81">J</emphasis>
Fruit
<emphasis bold="true" pageId="12" pageNumber="81">K</emphasis>
Mericarp valve
<emphasis bold="true" pageId="12" pageNumber="81">L</emphasis>
Columella
<emphasis bold="true" pageId="12" pageNumber="81">M</emphasis>
Seed (ventral face). (Source:
<emphasis bold="true" pageId="12" pageNumber="81">A-G</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Calatayud et al. 4711</emphasis>
, MO;
<emphasis bold="true" pageId="12" pageNumber="81">H-I</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Monteagudo et al. 4458</emphasis>
, US;
<emphasis bold="true" pageId="12" pageNumber="81">J</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">
<normalizedToken originalValue="Vásquez">Vasquez</normalizedToken>
&amp; Valenzuela 37638
</emphasis>
, MO;
<emphasis bold="true" pageId="12" pageNumber="81">K-M</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Monteagudo &amp; Ortiz 4605</emphasis>
, US).
</paragraph>
</caption>
<caption ID-Zenodo-Dep="995267" doi="10.3897/phytokeys.85.14757.figure2" httpUri="https://binary.pensoft.net/fig/153292" pageId="12" pageNumber="81" start="Figure 2" startId="F2">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Figure 2.</emphasis>
Distribution map of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron esseri" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="esseri">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron esseri</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<caption ID-Zenodo-Dep="995269" doi="10.3897/phytokeys.85.14757.figure3" httpUri="https://binary.pensoft.net/fig/153293" pageId="12" pageNumber="81" start="Figure 3" startId="F3">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Figure 3.</emphasis>
Morphology of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
(
<emphasis bold="true" pageId="12" pageNumber="81">A-J, L-M</emphasis>
) and
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
(
<emphasis bold="true" pageId="12" pageNumber="81">K, N</emphasis>
).
<emphasis bold="true" pageId="12" pageNumber="81">A</emphasis>
Habit, with paired branching and staminate inflorescences; note latex at damaged nodes
<emphasis bold="true" pageId="12" pageNumber="81">B</emphasis>
Leaf base (adaxial) with basal lobes
<emphasis bold="true" pageId="12" pageNumber="81">C</emphasis>
Leaf base (abaxial) with curled glandular margin
<emphasis bold="true" pageId="12" pageNumber="81">D</emphasis>
Staminate inflorescence with cymules subtended by glands; central flowers abscised leaving two lateral buds per cymule
<emphasis bold="true" pageId="12" pageNumber="81">E</emphasis>
Branch tip showing leaf-opposed inflorescence and stipule-enclosed renewal shoot
<emphasis bold="true" pageId="12" pageNumber="81">F</emphasis>
Summit of peduncle showing bract scars
<emphasis bold="true" pageId="12" pageNumber="81">G</emphasis>
Nearly mature green fruit
<emphasis bold="true" pageId="12" pageNumber="81">H</emphasis>
Mature brown fruit
<emphasis bold="true" pageId="12" pageNumber="81">I</emphasis>
Seed with funicle; holes in I &amp; J come from insect predation
<emphasis bold="true" pageId="12" pageNumber="81">J</emphasis>
Mericarp valve with outline of seed position
<emphasis bold="true" pageId="12" pageNumber="81">K</emphasis>
Mericarp valve with a seed; funicle obscures gap
<emphasis bold="true" pageId="12" pageNumber="81">L</emphasis>
Seed coat, transverse view (SEM)
<emphasis bold="true" pageId="12" pageNumber="81">M</emphasis>
Pericarp profile and top half of valve (exocarp removed)
<emphasis bold="true" pageId="12" pageNumber="81">N</emphasis>
Pericarp profile and top half of valve. (Abbreviations: f = funicle, g = gap, ib = inflorescence bract scar, p = pistillate, pb = pistillate bract scar, s = staminate. Orientation of
<emphasis bold="true" pageId="12" pageNumber="81">M-N</emphasis>
relative to
<emphasis bold="true" pageId="12" pageNumber="81">J-K</emphasis>
shown by diagrams where x-y = plane of cross section, z = apically pointing arrow. (Source:
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">Incadendron</taxonomicName>
,
<emphasis bold="true" pageId="12" pageNumber="81">A-E, G-H</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Farfan et al. 1049</emphasis>
,
<emphasis italics="true" pageId="12" pageNumber="81">1131</emphasis>
;
<emphasis bold="true" pageId="12" pageNumber="81">F</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Farfan et al. 706</emphasis>
, MO;
<emphasis bold="true" pageId="12" pageNumber="81">I-J, M</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Monteagudo &amp; Ortiz 4605</emphasis>
, US;
<emphasis bold="true" pageId="12" pageNumber="81">L</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Monteagudo et al. 4484</emphasis>
, US.
<taxonomicName authorityName="Steyerm" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis croizatii" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="croizatii">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis croizatii</emphasis>
</taxonomicName>
,
<emphasis bold="true" pageId="12" pageNumber="81">K, N</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Radosavljevic 296</emphasis>
, US).
</paragraph>
</caption>
<caption ID-Zenodo-Dep="995271" doi="10.3897/phytokeys.85.14757.figure4" httpUri="https://binary.pensoft.net/fig/153294" pageId="12" pageNumber="81" start="Figure 4" startId="F4">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Figure 4.</emphasis>
Micromorphological and anatomical features of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
.
<emphasis bold="true" pageId="12" pageNumber="81">A</emphasis>
Staminate flower with one anther removed, showing short filaments and basally connate sepals
<emphasis bold="true" pageId="12" pageNumber="81">B</emphasis>
Young inflorescence, showing inflorescence bract and staminate cymules subtended by glands and bracts
<emphasis bold="true" pageId="12" pageNumber="81">C</emphasis>
Pollen; mesocolpium-centered equatorial view of whole grain
<emphasis bold="true" pageId="12" pageNumber="81">D</emphasis>
Pollen; slightly oblique polar view of whole grains
<emphasis bold="true" pageId="12" pageNumber="81">E</emphasis>
Abaxial laminar surface with marginal gland
<emphasis bold="true" pageId="12" pageNumber="81">F</emphasis>
Abaxial laminar surface closeup, showing striate micro-sculpturing and stomata
<emphasis bold="true" pageId="12" pageNumber="81">G</emphasis>
Fractured shoot tip, showing stipule surrounding young leaf with conduplicate ptyxis
<emphasis bold="true" pageId="12" pageNumber="81">H</emphasis>
Anatomical cross section of shoot apex, showing central terminal inflorescence surrounded by nested series of two developing leaves with subtending sheathing stipules (composite tiled image)
<emphasis bold="true" pageId="12" pageNumber="81">I</emphasis>
Closer view of anatomical cross section of stipule, showing mucilage-secreting cell layer. (Abbreviations: a = site of attachment of missing anther, cb = cymule bract, g = glands, ib = inflorescence bract, in = inflorescence, lf = leaf, mu = dried mucilage, se = secreting cells, st = stipule.
<emphasis bold="true" pageId="12" pageNumber="81">A-G</emphasis>
imaged with a Zeiss EVO MA15 SEM at 10-12 kV after sputter coating with 25 nm of Au/Pd; SEM samples untreated and directly mounted from dried herbarium specimens; pollen from dehiscing anthers.
<emphasis bold="true" pageId="12" pageNumber="81">H-I</emphasis>
from paraffin-embedded, rehydrated herbarium specimens; 7
<normalizedToken originalValue="μm">μm</normalizedToken>
sections stained with iron-mordanted safranin O and celestine blue B; imaged with a Zeiss Universal. Source:
<emphasis bold="true" pageId="12" pageNumber="81">A, C-D</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Monteagudo et al. 4458</emphasis>
, US;
<emphasis bold="true" pageId="12" pageNumber="81">B</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Neill &amp; Kajekai 16622</emphasis>
, US;
<emphasis bold="true" pageId="12" pageNumber="81">E-I</emphasis>
<emphasis italics="true" pageId="12" pageNumber="81">Neill &amp; Kajekai 16646</emphasis>
, US).
</paragraph>
</caption>
<caption ID-Table-UUID="9FED5A1DB7703AE03E76D8FEAB57F42A" httpUri="http://table.plazi.org/id/9FED5A1DB7703AE03E76D8FEAB57F42A" pageId="12" pageNumber="81" start="Table 1" startId="T1">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Table 1.</emphasis>
Comparison of
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
with the morphologically most-similar neotropical genera. Based on primary observations with supplements from
<bibRefCitation author="Kruijt, RC" journalOrPublisher="Bibliotheca Botanica" pageId="14" pageNumber="83" pagination="1 - 109" refId="B15" refString="Kruijt, RC, 1996. A taxonomic monograph of Sapium Jacq., Anomostachys (Baill.) Hurus., Duvigneaudia J. Leonard and Sclerocroton Hochst. (Euphorbiaceae tribe Hippomaneae. Bibliotheca Botanica 146: 1 - 109" title="A taxonomic monograph of Sapium Jacq., Anomostachys (Baill.) Hurus., Duvigneaudia J. Leonard and Sclerocroton Hochst. (Euphorbiaceae tribe Hippomaneae." volume="146" year="1996">Kruijt (1996)</bibRefCitation>
and
<bibRefCitation author="Esser, H-J" journalOrPublisher="Mitteilungen aus dem Institut fuer Allgemeine Botanik Hamburg" pageId="13" pageNumber="82" pagination="121 - 133" refId="B6" refString="Esser, H-J, 1995. A taxonomic revision of Senefelderopsis Steyerm. (Euphorbiaceae), including additional notes on Dendrothrix Esser. Mitteilungen aus dem Institut fuer Allgemeine Botanik Hamburg 25: 121 - 133" title="A taxonomic revision of Senefelderopsis Steyerm. (Euphorbiaceae), including additional notes on Dendrothrix Esser." volume="25" year="1995">Esser (1995</bibRefCitation>
,
<bibRefCitation author="Esser, H-J" editor="Radcliffe-Smith, A" journalOrPublisher="Royal Botanic Gardens, Kew, UK" pageId="13" pageNumber="82" pagination="352 - 98" refId="B7" refString="Esser, H-J, 2001. Tribe 49. Hippomaneae. In: Radcliffe-Smith, A, Ed., Genera Euphorbiacearum. Royal Botanic Gardens, Kew, UK: 352 - 98" title="Tribe 49. Hippomaneae." volumeTitle="Genera Euphorbiacearum." year="2001">2001</bibRefCitation>
). The circumscription of
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sebastiania" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Sebastiania" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Sebastiania</emphasis>
</taxonomicName>
is controversial leading to some uncertainty in the breadth of character states.
</paragraph>
</caption>
<paragraph pageId="12" pageNumber="81">
<table pageId="12" pageNumber="81">
<tr pageId="12" pageNumber="81">
<th colspan="1" pageId="12" pageNumber="81" rowspan="1">Character</th>
<th colspan="1" pageId="12" pageNumber="81" rowspan="1">
<taxonomicName authorityName="K. Wurdack &amp; Farfan" authorityYear="2017" class="Magnoliopsida" family="Euphorbiaceae" genus="Incadendron" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Incadendron" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Incadendron</emphasis>
</taxonomicName>
</th>
<th colspan="1" pageId="12" pageNumber="81" rowspan="1">
<taxonomicName authorityName="Steyermark" authorityYear="1951" class="Magnoliopsida" family="Euphorbiaceae" genus="Senefelderopsis" higherTaxonomySource="CoL" kingdom="Plantae" lsidName="Senefelderopsis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Senefelderopsis</emphasis>
</taxonomicName>
</th>
<th colspan="1" pageId="12" pageNumber="81" rowspan="1">
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sapium" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Sapium" order="Malpighiales" pageId="12" pageNumber="81" phylum="Angiospermae" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Sapium</emphasis>
</taxonomicName>
</th>
<th colspan="1" pageId="12" pageNumber="81" rowspan="1">
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sebastiania" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Sebastiania" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="12" pageNumber="81">Sebastiania</emphasis>
</taxonomicName>
</th>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Stipules</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">
Lanceolate, sheathing, 10-13
<normalizedToken originalValue="×">x</normalizedToken>
3-4 mm, deciduous leaving elliptic scar
</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">
Lanceolate, 2.5-5
<normalizedToken originalValue="×">x</normalizedToken>
0.5-0.8 mm, deciduous leaving trigonous scar
</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">
Triangular, usually &lt;2
<normalizedToken originalValue="×">x</normalizedToken>
2 mm but up to 8 mm long, persistent (rarely tardily deciduous)
</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Lanceolate, small, persistent</td>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Indument</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Absent</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Present; multicellular, uniseriate</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Absent</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">
Usually absent; multicellular, uniseriate in some potential segregates (e.g.,
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Sebastiania" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Sebastiania vestita" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="vestita">
<emphasis italics="true" pageId="12" pageNumber="81">Sebastiania vestita</emphasis>
</taxonomicName>
<normalizedToken originalValue="Müll">Muell</normalizedToken>
. Arg.)
</td>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Leaf features</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Coriaceous; glands along margin; margin entire; ptyxis conduplicate</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Coriaceous; large glands at base and rimmed laminar glands; margin entire; ptyxis supervolute-curved</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Usually coriaceous; petiolar glands 2 (0 or 4), laminar glands absent; margin entire, glandular or toothed; ptyxis involute</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Membranous to subcoriaceous; glands absent or rimmed glands near base; margins usually minutely toothed; ptyxis involute</td>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Inflorescence structure</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Terminal, simple thyrse; 0-1 pistillate proximal, staminate distal in numerous 3-flowered cymules; cymule bract glands 4-5 per side, discoid; bracteoles absent</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Terminal, compound thyrse; 0-2 pistillate proximal per branch, staminate distal in numerous 5-10 flowered cymules; cymule bract glands 1 per side, fleshy and elongate; bracteoles present</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Terminal or axillary, simple thyrse; 0-15 pistillate proximal, staminate distal in numerous 2-16 flowered cymules; cymule bract glands 1 per side, elliptic, flattened; bracteoles present</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Terminal or axillary, simple thyrse; 0-5 pistillate proximal, staminate distal in numerous 1-3(7)-flowered cymules; cymule bract with 1 discoid gland to large segmented glandular mass per side; bracteoles usually absent</td>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Staminate flowers</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Stamens 3; sepals 3, connate at base</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Stamens 3-5; sepals 3, connate at base</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Stamens 2; sepals 2, connate at base to 2/3 of length</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Stamens 3; sepals 3, connate at base</td>
</tr>
<tr pageId="12" pageNumber="81">
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Fruits</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Large; pericarp thick, woody</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Large; pericarp thick, woody</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Small to large; pericarp woody to leathery, usually thin</td>
<td colspan="1" pageId="12" pageNumber="81" rowspan="1">Small to large; pericarp thin</td>
</tr>
</table>
</paragraph>
<caption ID-Zenodo-Dep="995273" doi="10.3897/phytokeys.85.14757.figure5" httpUri="https://binary.pensoft.net/fig/153295" pageId="12" pageNumber="81" start="Figure 5" startId="F5">
<paragraph pageId="12" pageNumber="81">
<emphasis bold="true" pageId="12" pageNumber="81">Figure 5.</emphasis>
Hippomaneae shoot tips showing stipule and ptyxis variation.
<emphasis bold="true" pageId="12" pageNumber="81">A</emphasis>
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Homalanthus" higherTaxonomySource="GBIF,CoL" kingdom="Plantae" lsidName="Homalanthus nutans" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="nutans">
<emphasis italics="true" pageId="12" pageNumber="81">Homalanthus nutans</emphasis>
</taxonomicName>
(G. Forst.) Guill. with large, deciduous, sheathing stipules protecting shoot tip
<emphasis bold="true" pageId="12" pageNumber="81">B</emphasis>
Transverse view of
<taxonomicName lsidName="H. nutans" pageId="12" pageNumber="81" rank="species" species="nutans">
<emphasis italics="true" pageId="12" pageNumber="81">H. nutans</emphasis>
</taxonomicName>
shoot tip, showing nested series of three developing leaves with scroll-like lamina (ptyxis involute), each surrounded by a pair of sheathing stipules
<emphasis bold="true" pageId="12" pageNumber="81">C</emphasis>
Transverse view of
<taxonomicName class="Magnoliopsida" family="Euphorbiaceae" genus="Excoecaria" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Excoecaria cochinchinensis" order="Malpighiales" pageId="12" pageNumber="81" phylum="Tracheophyta" rank="species" species="cochinchinensis">
<emphasis italics="true" pageId="12" pageNumber="81">Excoecaria cochinchinensis</emphasis>
</taxonomicName>
Lour. young leaf showing slightly inrolled lamina (ptyxis conduplicate-involute)
<emphasis bold="true" pageId="12" pageNumber="81">D</emphasis>
<taxonomicName lsidName="E. cochinchinensis" pageId="12" pageNumber="81" rank="species" species="cochinchinensis">
<emphasis italics="true" pageId="12" pageNumber="81">E. cochinchinensis</emphasis>
</taxonomicName>
shoot tip with small persistent stipules. (Abbreviation: s = stipule. Source: Freehand sections of fresh tissues grown in Department of Botany greenhouses).
</paragraph>
</caption>
</subSubSection>
</treatment>
</subSection>
</document>