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Notes on Justicia sect. Monechma (Acanthaceae) in Angola, including two new species

Authors:
Iain Darbyshire at Royal Botanic Gardens, Kew
Iain Darbyshire
David Goyder at Royal Botanic Gardens, Kew
David Goyder
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Abstract and Figures

Two new species of Justicia sect. Monechma, Justicia cubangensis and J. eriniae, are described from Angola. Their affinities are reviewed, together with a discussion on the correct generic name to apply to these species. A conspectus of Justicia sect. Monechma ‘Group I’ in Angola is presented, including a key to the species and notes on habitat and distribution for each taxon. Two new combinations and one new name are provided for species for which there was no previous name available in Justicia, and lectotypes are proposed for four names. Twelve species are recognised in this group in total, eight of which are currently believed to be endemic to Angola. Justicia subsessilis is recorded in Angola for the first time, and J. laeta is placed in sect. Monechma, having previously been tentatively referred to sect. Tyloglossa. Monechma carrissoi is considered to be conspecific with J. virgultorum and so is synonymised here. In addition, the botanical importance of the type locality of J. cubangensis, the Cuchi River gorge of the Cubango drainage, is highlighted and two further new provincial records for Cuando Cubango are noted from this site: Eriocaulon angustibracteum (Eriocaulaceae) and Crepidorhopalon schweinfurthii (Linderniaceae).
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Blumea 64, 2019: 97–107
www.ingentaconnect.com/content/nhn/blumea https://doi.org/10.3767/blumea.2019.64.02.01
RESEARCH ARTICLE
INTRODUCTION
Angola remains very unevenly explored botanically despite
its rich and diverse flora (Romeiras et al. 2014, Goyder &
Gonçalves2019).Followingtheendoftheprolongedcivilwar
(1975–2002)andthesubsequentgradualincreaseinacces-
sibilityofmostofthecountrytoscientificstudy,therehasbeen
arenewed interestinthebotanyofAngola.Thiswasfuelled
by the publication of the checklist of vascular plants of Angola
(Figueiredo&Smith2008),andhascontinuedthroughaseries
of biodiversity surveys in under-explored parts of the countries
(e.g.,seeDarbyshireetal.2014,Gonçalves&Goyder2016,
Goyder&Gonçalves2019)whichhaveconsistentlyresulted
inthediscoveryofasignificantnumberofnewplantspecies
to science and/ ornewplantrecordsforAngola.Inthispaper,
we report on the discovery of two new species of Justicia L.
sect.Monechma (Hochst.)T.Anderson,foundduringtwosuch
recent expeditions, and we provide a synoptic review of this
groupinAngola.
ThefirstofthenewspeciesofJusticia was discovered in May
2015 as part of a major biodiversity assessment of the upper
catchment of the Okavango system in Angola – the Okavango
Wilderness Project – during which the second author had the
opportunitytovisittheCuchiRivergorge.The17specimens
collected at this site included a perennial species of Justicia
sect.Monechma, a single plant of which was found growing
amongstgrassinopenmiombowoodlandclosetothegorge.
OnstudyingthespecimenatKewherbarium(K)andcompar-
ingittotypematerialavailableonJSTORGlobalPlant(https://
plants.jstor.org/),thespecieswasfoundtobemostsimilarto
Justicia lolioides S.Moore(1880:310)(=Monechma lolioides
(S.Moore) C.B.Clarke) from the Pungo Andongo region of
northern Angola, but differing in a number of key characters
(seeTable1).AsubsequentreviewofspecimensintheBM,K
andLISCherbaria,whichtogetherhousethelargesthistorical
collections of Angolan plants, revealed three earlier collections
of the same species from the Menongue and Kuvango areas
(Gossweiler 2701, 3100, Santos & Barroso 2674).TheGoss-
weiler specimens at BM had been misnamed as M. lolioides
byMoore(1930)andcontinuedtobefiledunderthatnameto
thisday,whiletheSantos&Barrosospecimenwasonlynamed
togenus.These four collectionstogether form the basisfor
the new species, Justicia cubangensis I.Darbysh.& Goyder,
describedbelow.
ThesecondnewlydescribedspecieswasdiscoveredinApril
2017 during Acanthaceae-focussedfieldresearchinNamibe
ProvinceofSouthwestAngolabyErinTrippoftheUniversityof
Colorado(COLOherbarium)andKyleDexteroftheUniversity
ofEdinburgh.Thisfieldworkwascarriedoutprimarilytostudy
the genus Petalidium Nees in Angola, where species are con-
centrated in the Kaokoveld Centre of floristic endemism which
extends from northwest Namibia into the arid southwestern
portionofAngola(VanWyk&Smith2001,Trippetal.2017).
Theopportunitywasalsotakentocollectspecimensofother
Acanthaceaespeciesas they were encountered.Thesecol-
lections have revealed several new or interesting species,
including Barleria deserticola I.Darbysh.& E.Tripp, currently
underdescription(Darbyshireet al. 2019), and aspeciesin
the tribe Ruellieae which is unmatched and yet to be placed to
genus(E.A.Tripp,unpubl.data).Aduplicateofthespecimen
of a perennial species of Justicia sect.Monechma, collected
between Bibala and Assunçao (Tripp & Dexter 6917), was
senttothefirstauthoratK.Thespecimenwasreceivedjust
as a key to the Angolan species of Justiciasect.Monechma
was in preparation for the current work, and it was immediately
recognisedasanewspeciesforAngola.Oncheckingtheac-
countsof the genusforsouthernAfricaandNamibia(Meyer
1968,Badenetal.1995)andthe‘FloraofTropicalEastAfrica’
and‘FloraZambesiaca’regions(Vollesen2010,2015),itwas
clear that this is a new species to science, apparently restricted
Notes on Justicia sect. Monechma (Acanthaceae) in Angola,
including two new species
I.Darbyshire1,*,D.J.Goyder1,2
1 RoyalBotanicGardens,Kew,Richmond,Surrey,TW93AB,UnitedKing-
dom; corresponding author e-mail: i.darbyshire@kew.org.
2 NationalGeographicOkavangoWildernessProject,WildBirdTrust,South
Africa.
Key words
Cuando Cubango
Huíla
justicioid
miombo
Namibe
taxonomy
Abstract TwonewspeciesofJusticiasect.Monechma, Justicia cubangensis and J. eriniae, are described from
Angola.Theiraffinitiesarereviewed,togetherwithadiscussiononthecorrectgenericnametoapplytothesespecies.
A conspectus of Justicia sect.Monechma ‘GroupI’inAngolaispresented,includingakeytothespeciesandnotes
onhabitatanddistributionforeachtaxon.Twonewcombinationsandonenewnameareprovidedforspeciesfor
which there was no previous name available in Justicia,andlectotypesareproposedforfournames. Twelvespecies
arerecognisedinthisgroupintotal,eightofwhicharecurrentlybelievedtobeendemictoAngola.Justicia sub-
sessilis isrecordedinAngolaforthefirsttime,andJ. laeta isplacedinsect.Monechma, having previously been
tentativelyreferredtosect.Tyloglossa. Monechma carrissoiisconsideredtobeconspecificwithJ. virgultorum and
soissynonymisedhere.Inaddition,thebotanicalimportanceofthetypelocalityofJ. cubangensis,theCuchiRiver
gorge of the Cubango drainage, is highlighted and two further new provincial records for Cuando Cubango are noted
from this site: Eriocaulon angustibracteum(Eriocaulaceae)andCrepidorhopalon schweinfurthii(Linderniaceae).
Published on3May2019
98 Blumea–Volume64/2,2019
tothenorthernNamibDesertofAngolaandunmatchedinthe
herbariumcollectionsstudied.ItisdescribedbelowasJusticia
eriniaeI.Darbysh.
ThisworkformsapartofwiderstudiesontheAcanthaceae of
Angola, focussing on the diverse genera Barleria L.(Darbyshire
etal.2019),Justicia and Petalidium(seeTrippetal.2017),but
with the eventual aim of documenting all the genera of Acan-
thaceaethere.RecentstudiesonothergeneraofAcanthaceae,
including Blepharis Juss.(Vollesen2000),Justiciasect.Harni-
eria Benth.(Hedrén1989),Phaulopsis Willd.(Manktelow1996),
Hypoestes R.Br.(Darbyshire 2015)andRhinacanthus Nees
(Darbyshireetal.2018),haveallresultedinthedescriptionof
new,endemicspeciesinAngola.
WHAT IS THE CORRECT GENERIC NAME TO APPLY TO
THE NEW SPECIES – MONECHMA OR JUSTICIA?
Monechma Hochst. has traditionally been separated from
Justicia based on the presence of 2-seeded capsules with
compressed seeds with a smooth testa, vs usually 4-seeded
capsules with seeds variously sculptured or if smooth then not
socompressed(Hedrén1990,Badenetal. 1995).However,
Hedrén(1990)foundthathisnewspeciesJusticia tetrasperma
Hedrén has a combination of characters intermediate between
Justicia and Monechma, namely that it has a 4-seeded capsule
withcompressedsmoothseeds.This,togetherwithhisobser-
vationthatitcanbedifficulttoseparate membersofthetwo
generaintheabsenceofmaturefruits,ledHedréntosuggest
that Monechma is better treated as a section of Justicia as per
Anderson(1863).Thisdecisionhasbeenfollowedbyseveral
subsequentauthors(e.g.,Darbyshireetal.2010,2015b,Man-
ning&Goldblatt2014),althoughothershavechosentomain-
tain Monechma asadistinctgenus(e.g.,Baden etal.1995,
Ensermu2006,Makholela2008).However,recentmolecular
phylogeneticstudiesonthe‘justicioid’lineage(Kieletal.2017)
indicate that Justicia sect.Monechma is not monophyletic, with
two distinct and widely separated lineages discernible based
onlimitedsampling(sixspeciesofsect.Monechma sampled).
Monechma GroupIofKieletal.(2017),isbelievedtocomprise
predominantly tropical African species including the type spe-
cies of the genus, M. bracteatum Hochst.(=Justicia bracteata
(Hochst.) Zarb), and falls within their ‘Core Harnieria’ clade
together with members of Justicia sect.Harnieria. Monechma
GroupIIofKieletal.(2017),isbelievedtocomprisepredomi-
nantly southern African species and falls within the Diclipterinae
lineage, sister to the genus Rhinacanthus Nees.
Although these two groups of Monechma are morphologically
similar, particularly in flower and fruit morphology, there is some
morphologicalevidencetosupporttheirseparation.Inparticular,
therearenotabledifferencesininflorescenceform.InGroupI
the flowers are usually arranged in 1–few-flowered cymes ag-
gregated into axillary and/or terminal spikes or fascicles with
thebractsmarkedlydifferentiatedfromtheleaves.Theseplants
are annual or perennial herbs or suffrutices often growing in
fire-pronewoodedgrasslands,althoughannualspeciessuch
as J. bracteata canbeweedsofdisturbedareas.InGroupII,
thespecieshavesingle-orrarely2-flowered(sub)sessile axil-
lary inflorescences which can together sometimes form weakly
definedterminalspikesbutwiththebractslargelyundifferenti-
atedfromtheleaves.SpeciesofGroupIIaremainlyfoundin
arid environments and often have either highly reduced and/or
densely glandular-hairy leaves, while most are small shrublets
(Kielet al. 2017). Based on these diagnosticmorphological
characters, the two new species described here are hypothe-
sised to belong within Monechma GroupI.
Kieletal.(2017)reviewedthepotentialtaxonomicimplications
oftheirfindingsofthecomplexrelationshipsbetweenandwithin
generainthe‘justicioid’lineage.They noted thatinorderto
maintain a broadly circumscribed Justicia, including morphologi-
cally similar taxa such as Monechma s.lat.,theentire‘justicioid’
lineagewouldpotentiallyhavetobetreatedasasinglegenus.
Thisisundesirableasitwouldmeansubsumingseverallarge
genera that are easily separated morphologically, such as Di-
cliptera Juss. and Hypoestes R.Br.Ifacceptingaparaphyletic
Justicia is not an option, the most credible alternative appears
to be to divide Justicia s.lat. intoanumberofsegregategenera.
Underthisscenario,the‘CoreHarnieria’ clade could be consi-
dered a separate genus, with Monechma the earliest available
name.ThegenusMonechma could therefore be resurrected,
albeit with a revised circumscription, and the new species de-
scribed here would be renamed in Monechma.However,Kiel
etal.(2017)notethatonly12–15 % of all justicioids have been
sampledsofarandanumberofsamplingdeficienciesneedto
be addressed before fully informed taxonomic decisions can be
made.ThisiscertainlytrueofbothgroupsofMonechma where
only a small number of species were sampled, and an incom-
plete range of the morphological variation within Monechma
s.lat.wasrepresented.Inviewofthisuncertainty,weconsider
it best to maintain Justicia sect.Monechma at the present time,
and the new species are described in Justicia.
MATERIALS AND METHODS
Thisstudyisbasedupontheinvestigationofherbariumspeci-
mensheldatBM,COLO,K,LISC and LUBAherbaria,sup-
ported where available by photographs of the species in the
field.Abbreviationsforherbariafollowthestandardformlisted
onIndex Herbariorum (Thiers, continuously updated). The
fledglingherbariumoftheAngolanMinistryoftheEnvironment’s
InstitutoNacionaldaBiodiversidadeeÁreasdeConservação,
Character Justicia cubangensis Justicia lolioides
Indumentumofinorescencerachis,bracts, Puberulouswithmixedeglandularand Glabrousexceptforshortascendingeglandularhairsalong
bracteoles and calyces capitate-glandular hairs margins of bracts, bracteoles and calyx lobes; calyces can
have sessile glands
Bract, bracteole and calyx colouration Apices dark green or purplish, margins with a Apices darker only along midrib, margins with a broad pale
narrow pale portion portion
Bractlength:mid-portionofspike 1.8– 4.8mm,markedlyshorterthancalyces 6 –7.5mm,equaltooronlyslightlyshorterthancalyces
Bracteolelength 2.5–5.5mm,markedlyshorterthancalyces 7– 8.5mm,±equalinlengthtocalyces
Corolla indumentum: lower lip Finely eglandular-puberulous Mixed eglandular-puberulous and numerous coarser
conspicuous bristly eglandular hairs
Antherappendage Apextaperingtoasinglepoint,notbifid Apexbifid
Capsuleindumentum Eglandular-puberulous Glabrousexceptoccasionallyoneortwohairsonshortbeak
Capsulelength ±11mm(immature) 8–9mm
Table 1 Morphological characters separating Justicia cubangensis from J. lolioides.
99
I.Darbyshire&D.J.Goyder:Justicia section Monechma in Angola
whichisnotyetlistedon IndexHerbarium,isabbreviatedto
‘INBAC’.All duplicates of specimenslisted havebeen seen
exceptwherenotedas‘[notseen]’.Thoseforwhichonlyonline
imageshavebeenseenaremarkedwithanasterisk(*);these
wereaccessed viaJSTORGlobalPlants (https://plants.jstor.
org/).Allmeasurementsweremadeondriedmaterialexcept
forthoseoftheflowers,whichweresoakedinAerosolOT5%
solutionpriortodissectionandmeasurement.
SpellingofAngolanprovincesandmajortownsorcitiesfollows
usageinFigueiredo&Smith(2008)exceptforthemunicipality
of Cubango which is now generally referred to as Kuvango, and
theprovinceCuandoCubangowherethehyphen isomitted.
Historicplacenamesorotherspellingvariantsareplacedwithin
squarebracketsafterthestandardisedname.
Theassessmentof the conservation statusofthe new spe-
ciesfollowsthecategoriesandcriteriaoftheIUCNRedListof
ThreatenedSpecies(IUCN2012).
Fig. 1 Justicia cubangensis I.Darbysh.&Goyder.a.Habit,basalanddistalportionsofstems;b.matureleaf,abaxialsurface;c.inflorescence;d.dissected
calyx,externalsurface,withdetailofindumentum;e.dissectedcorollawithandroecium;f.anther;g.pistil;h.immaturecapsule(Goyder 8068,CAS,K,LUBA).
—Scalebars:a=3cm;b– c=1cm;d – e,g – h=5mm;f=2mm.—DrawnbyJulietWilliamson.
c
b
da
g
e
f
h
100 Blumea–Volume64/2,2019
TAXONOMIC ACCOUNT: NEW SPECIES DESCRIPTIONS
Justicia cubangensis I.Darbysh.& Goyder, sp. nov. — Fig.
1, 2a – c; Map 1
Most similar to Justicia lolioides S.Moorebutdiffersintheinflorescenceha-
ving a puberulous indumentum of mixed eglandular and capitate-glandular
hairsthroughout(vsinJ. lolioides inflorescences largely glabrous except for
short ascending eglandular hairs along the margins of the bracts, bracteoles
andcalyxlobes);thecalyxlobesbeingmarkedlylongerthanthesubtending
bractsandbracteoles(vscalyx±equalinlengthtothebractsandparticularly
tothebracteoles);thecorollabeingfinelyeglandular-puberulousexternally
(vscorollahavingnumerouscoarsebristlyhairsonthelowerlipexternally
inadditiontobeingeglandular-puberulous);theantherappendagetapering
toasinglepointattheapex(vs antherappendage bifidattheapex);and
thecapsulebeingeglandular-puberulousand±11mmlong(vscapsulees-
sentiallyglabrousand8 – 9mm long).SeeTable1.—Type:Goyder 8068
(holoK;isoCAS,INBAC,LUBA),Angola,CuandoCubangoProvince,Cuchi
Rivergorge(Cubangodrainage),c.7kmNofCuchi,S14.58972E16.90722,
1350m,fl.&imm.fr.,28May2015.
Monechma lolioides sensuMoore(1930)138,non(S.Moore)C.B.Clarke
(1900)218.
Etymology.Theepithet‘cubangensis’denotestheCubangoRiverdrain-
ageinwhichthisspeciesisfound.
Erectordecumbentperennialherb,severalannual,many-branch-
ed stems 15–65 cm tall from a woody base and rootstock, burnt
stembasesfrompreviousyears’growthsometimespersisting;
stemsquadrangular,twooppositesideswithacentralfurrow,
other two opposite sides either flat or with two shallow furrows,
minutely retrorse-puberulous, hairs often more numerous on the
two centrally furrowed sides, most conspicuous on lower most
internodes;nodallinewithlongerhairs.Leaves sessile, linear-
lanceolate,longest blades 29 – 46 by 2.3–3.2 mm (length:
widthratio10 – 16:1);basecuneate,marginentire,apexacute,
surfaces with sparse and inconspicuous minute antrorse hairs
mainlyalongmargin;venationof5parallelveins,±prominent
particularlyonadaxialsurface.Inflorescences terminal and in
upperaxils, secund spikes 25–70 mm long; rachis(at least
distally),bracts,bracteolesandcalycespuberulouswithmixed
patent eglandular and capitate-glandular hairs; peduncle
3.5–12mmlong;eachnodeofinflorescencesingle-flowered;
bractspaired, minute, lanceolate or triangular, 1.8 – 4.8 by
0.7–1.5mmincentralportionofspike,darkgreenorpurplish
with paler base, midrib prominent; flowers sessile along rachis
or lowermost flowers on a secondary peduncle to 5 mm long;
calyx immediately subtended by erect paired bracteoles, lan-
ceolate,2.5–5.5by0.7–1.8mm,paleyellowishcreamdistally,
greentowardsapex,withpalehyalinemargin.Calyx divided
almosttobaseintofivesubequallinear-lanceolatelobes6–7.5
by0.7–1mminflower,extendingsomewhatin fruitwhenup
to9.5by1.6mm,paleyellowishcreamproximallyandalong
Fig. 2SpeciesofJusticia sect.Monechma fromAngola.a–c.Justicia cubangensis I.Darbysh.&Goyder;d–e.Justicia mooreiI.Darbysh.&Goyder;f.Justicia
virgultorum(S.Moore)I.Darbysh.&Goyder(a– c. Goyder 8068;d– e.Goyder 8210;f.Goyder 8471).—PhotosbyD.Goyder.
101
I.Darbyshire&D.J.Goyder:Justicia section Monechma in Angola
margin, dark green to pinkish brown towards apex and along
threeparallelveins.Corolla 12–13mmlong,cream-white,pal-
ateoflowerlippinkwithwhiteveinsinraised‘herring-bone’pat-
tern,eglandular-puberulousexternallymainlyonlips;tube±7.5
mmlong,cylindricalwithventricosefloor±2mmfrombase,
pubescent internally at base of pouch and at base of rugula
(stylarfurrow);upperliphooded,5.5– 6mmlong,apexshallowly
bilobed;lowerlip5.2–6.2mmlong,reflexeddistallyatanthesis,
apex3-lobed,laterallobes1.2–1.5by1–1.1mm,medianlobe
broader,1.7–1.8mmwide.Stamens inserted±5.5mmfrom
baseofcorollatube;filamentswhite,5.5–5.7mmlong;anthers
atfirstgreen,turningdarkbrown,withthecaeoverlappingfor
±halftheirlength,oblique,uppertheca1–1.2mmlong,lower
theca1.2–1.4mmlong,withbasalwhitesubulateappendage
1–1.3mmlong,boththecaewithfewirregularroundedprotru-
sions(?glands)oninnersurface.Ovary oblong-ovoid,1.4–1.7
mm long, sparsely puberulous towards apex; style 9–10 mm
long,pubescentinproximalhalf;stigmashortlybifid.Capsule
onlyseeninimmaturestate,2-seeded,±11mmlong,shortly
stipitate, eglandular-puberulous externally; immature seeds
somewhatcompressed,withsmoothsurface,glabrous.
 Distribution& Ecology—Justicia cubangensis is found
in Cuando Cubango and Huíla provinces of South-central
Angola.Itgrowsinopen, sunnyareasamongstgrasslandin
open miombo woodland and open thicket on sandy soils, at
±1350–1400melevation.
Conservation—This species is currently known only
fromthreelocalities:theCuchiRivergorge,thevicinityofthe
municipalityofMenongueandbetweenKuvangoandChipindo.
Onlyasingle plant was observedatCuchi River gorgeand
GossweilerrecordeditasrareatMenongue,whileSantos&
Barroso recorded it as occurring in scattered tufts near Kuvan-
go.ThisregionofAngolahasnotbeenthoroughlybotanized,
and furthermore, many Angolan Acanthaceae collections were
lost–presumeddestroyed–duringthecivilwar(seeDarbyshire
etal. 2019). It may therefore prove to be more widespread
and/ orlocallycommonthancurrent evidencesuggests.The
sites around Menongue are likely to have been impacted by
the continued development of this town and the related expan-
sionofagricultureinthesurroundingarea.Thespeciesmaybe
tolerant to some degree of disturbance, however, as the Cuchi
Rivergorgesitewasopenandwaspossiblyclearedfromthe
surrounding miombo. In light of the incomplete evidence to
date, Justicia cubangensisisprovisionallyassessedasData
Deficient(DD).
Additional specimens seen. AngolA, Gossweiler 2701 (BM[BM001135166],
?K-seenote),Menongue[Munongue],fl.&immfr.,20April1906;Gossweiler
3100(BM[BM001135165],K),Ganguellas[Gunguellas],atMenongue[Mu-
nongue],fl.&immfr.,April1906;Santos & Barroso 2674(LISC*),Kuvango
[ArturdePaiva],naestradaparaChipindo,fl.,10May1969.
 Notes—AsnotedintheIntroduction,thisspecieshasprevi-
ously been confused with Justicia lolioides with which it shares
narrowfoliageandveryslender,secundinflorescencespikes.
However,itiseasilyseparatedbythecharacterslistedinthe
diagnosisandinTable1.
InadditiontoaduplicateofGossweiler 3100(incorrectlylabel-
led as 30100),thereisasecond,unnumberedGossweilercol-
lection of this species at K with no locality data or date; it may
be a duplicate of Gossweiler 2701.
TheCuchiRivergorge,thetypelocalityforthisspecies,isasite
ofconsiderablebotanicalimportance.Itisoneofthefewplaces
in the region where rock emerges from the overlying sand and
is the locus classicus of succulents such as Aloe metallica
Engl.&GilgandEuphorbia faucicolaL.C.Leach.Inadditionto
the new species of Justicia, the 17 collections made at Cuchi
RivergorgeinMay2015includedtwonewbotanicalrecordsfor
CuandoCubangoProvince,recordedbelow.Asubsequentvisit
to the site in November 2016 revealed a species of Crotalaria
L.thatisalsoprobablynewtoscience.
Eriocaulon angustibracteumKimp.(1994)329(Eriocaulaceae)
Goyder 8071(INBAC,K,LUBA,PRE),Angola,CuandoCubangoProvince,
CuchiRivergorgec.7kmNofCuchi, 28May2015.
ThisisonlythethirdrecordofthisspeciesfromAngola.Itwas
notrecordedbyFigueiredo&Smith(2008)intheirchecklistof
Angolanplants,butacollectionatKfromHuamboprovince,
nearUkuma[Cuma],madein1959(Damann in Herb. Hess
59/ 228)wasredeterminedasthisspeciesbySylviaPhillipsin
2016.ItwasalsocollectednearCapaia,LundaNorte,in2013
(Goyder et al. 7163).Thespeciesisquitewidelydistributedand
hasbeenrecordedfromsouthernTanzania,Zambia,theKa-
tangaregionofD.R.Congo,andCameroon(Phillips2010:65).
Crepidorhopalon schweinfurthii(Oliv.)Eb.Fisch. (1989) 443;
(1992)130(Linderniaceae)
Goyder 8060(INBAC,K,LUBA,PRE),Angola,CuandoCubangoProvince,
CuchiRivergorgec.7kmNofCuchi, 28May2015.
ThisspecieswaslistedbyGhazanfar(2008)underthename
Torenia schweinfurthiiOliv.(Scrophulariaceae),andwithinAn-
golaithasbeenrecordedfromBenguela,Huíla,Malangeand
Moxicoprovinces.Ithasawidebutscattereddistributionfrom
Malito South Sudan, and south toMozambiqueandAngola
(Darbyshireetal.2015a:329).
Justicia eriniae I.Darbysh.,sp. nov. — Fig.3;Map1
SimilartoJusticia tricostata Vollesenbutdiffersinthecapsulebeingpuberu-
lousandhavingblackseeds(vscapsuleglabrous,seedsmottledgreyand
black in J. tricostata);thebractsandbracteolesbeingshorter,thelatterbeing
4.5– 6mmlong,typically±halfthelengthofthecalyx(vs6.5–12.5mmlong
andonlyslightlyshorterthanorsubequaltothecalyces);theinflorescence
beinglaxthroughout(vstypicallydensethroughoutoratleastdistally);and
thestem and inflorescence indumentumbeing ± patent (vsindumentum
antrorse). InAngola,itismostlikelytobeconfusedwithJusticia glaucifolia
(S.Moore)I.Darbysh. & Goyder(seebelow)which shares the laxsecund
inflorescences with an eglandular indumentum, but J. eriniae differs in the
leaves being lanceolate with a markedly acute apex and a length : width ratio
4.5– 6.7:1(vsleavesobovatewitharoundedorobtuseapex,length:width
Map 1DistributionmapofJusticia cubangensis (●)andJ. eriniae (■).
102 Blumea–Volume64/2,2019
ratio2.5–2.6:1inJ. glaucifolia);thestem,inflorescencerachisandcalyces
etc.havingaspreading/patentindumentum(vsindumentumantrorse);and
inhavinglargerflowers,thecalyxlobes8 –9mmlong,thecorolla18 –18.5
mmlong (vs calyx lobes±6mm long, corolla ±14mmlong).See Table
2.—Type:Tripp & Dexter 6917(holoK; isoCOLO[notseen],LUBA[not
seen]),Angola,NamibeProv.,between Bibala andAssunçao, S14.79891
E13.22217,189m,fl.&fr.,10Apr.2017.
Etymology.Theepithet‘eriniae’honoursDr.ErinTrippoftheMuseumof
NaturalHistory(COLOHerbarium)attheUniversityofColorado,aleading
authority on Acanthaceae, great all-round naturalist and collector of the type
specimenofthisspecies.Dr.Trippalsokindlyprovidedextrafloralmeasure-
mentsfromtheCOLOisotype.
Brittle-stemmed herb, with many-branched stems to 50–75 cm
tall;stemsquadrangularorsomewhat6-angular,pale-pubes-
cent throughout, hairs patent or those immediately above the
nodes somewhat deflexed. Leaves with a short, ill-defined
petioleto4.5mmlong,bladelanceolatetonarrowlyso,longest
Fig. 3 Justicia eriniae I.Darbysh.a.Habit,distalportion ofstem; b.mature leaf,abaxial surface;c. inflorescence;d. bracts,bracteolesandbaseofcalyx;
e.dissectedcalyx,externalsurfacewithdetailofindumentum;f.flower,lateralview;g.anther;h.capsulewithpersistentcalyxandbracteoles;i.seed(Tripp
& Dexter 6917,K).—Scalebars:a –b=3cm;c – f,h=1cm;g,i=3mm.—DrawnbyJulietWilliamson.
c
b
d
a
g
hi
e
f
103
I.Darbyshire&D.J.Goyder:Justicia section Monechma in Angola
blades57–65by8.5–13mm(length:widthratio4.5–6.7:1);
base cuneate, margin entire, apex acute, surfaces sericeous-
puberulous when young, more sparse at maturity when hairs
most numerous along veins beneath, midrib above and margin;
lateralveins2or3perside,markedlyascending,withscalari-
formtertiaryvenationbetweenmidribandlateralveins.Inflo-
rescences terminal and sometimes also in upper axils, secund
spikes5.5 – 8 cm long,each node of inflorescence single-
flowered, lax, up to 9 nodes along spike; peduncle 10–19 mm
long, rachis eglandular-puberulous; bracts paired, lanceolate,
3.5–5.5by1.5mmincentralportionofspike(thoseatlower-
mostfertilenodecanbelongerandmorelinear),atfirstpaleyel-
lowishwith3prominent(brown-)greenveins,becomingbrown
with age; flowers sessile, calyx immediately subtended by erect
pairedbracteoles,resemblingbractsbut4.5–6by1– 2 mm,
apex±attenuate,3-veinedbutmidveinapparentlycomposedof
twopartiallyfusedparallelveins.Calyx divided almost to base
intofivesubequallinear-lanceolatelobes8– 9by1.3– 2mmin
flower,extendingsomewhatinfruitwhenupto10.5mmlong,
pale yellowish cream with three markedly raised parallel green
veins, apices green, external surface eglandular-puberulous,
hairsalongthemarginslongerandsomewhatcrisped.Corolla
18–18.5mmlong,white,eglandular-pubescentexternally;tube
±10.5mmlongincludingunexpandedbasalportion2.5–3.5
mmlong, floor of expanded throat markedly ventricose,6.5
mm deep centrally (when flattened), 3 mm deep at mouth;
upperliphooded,±8mmlong,apexshallowlybilobed;lower
lip±8.5mmlong,apex3-lobed,lobesrounded,2.5mmlong,
median lobe broader, palate of lower lip with weakly raised
‘herring-bone’pattern.Stamens inserted±7–8mmfrombase
ofcorollatube;filaments6–7.5mmlong;antherswiththecae
overlappingfor ± 1/3 theirlength, somewhat oblique,upper
theca1.2mmlong,lowertheca1.45–1.5mmlong,withbasal
paleyellowish appendage 1.7–1.9mm long, apex unevenly
andinconspicuouslyforked,antherconnectivepiloseatapex.
Ovary oblong-ovoid,±3mmlong,sparselypuberuloustowards
apex;style±16mmlong,sparselypuberulous,hairsbecom-
ingdenserinproximalportion;stigmaminutelybifid,onelobe
muchreduced.Capsule 2-seeded,10.5 –11mmlong,shortly
stipitate, eglandular-puberulous externally; seeds lenticular,
compressed,black,3.5–4mmdiam,withsmoothsurface.
 Distribution&Ecology—Justicia eriniae is recorded from
NamibeProvinceofSouthwestAngolawhereitisknownonly
fromthetypespecimen.Itwasrecordedfromroadsidethorny
scrub in lowland Colophospermum mopane woodlandat189
melevation.
Conservation — Only a small population of fewer than 10
plants was recorded at the single locality known for this spe-
cies, but the surrounding areas were not extensively searched
by the collectors, and potentially suitable habitat is widespread
intheregion.Althoughitwascollectedfromaroadsidesite,it
wasinalargelyundisturbedareawithverylittletraffic.Mining
of marble occurs in the vicinity of the type locality but does not
appeartoimpactthisspecies(E.A.Tripp,pers.comm.).With
more information needed on its full range and abundance,
J. eriniae iscurrentlyconsideredtobeDataDeficient(DD)but
it may prove to be of Least Concern as threats appear to be
minimal.
 Taxonomicnotes—Justicia eriniae is most likely to be
confused with J. tricostata, a recently described species from
rockoutcropsandstonysoilsinmiombowoodlandinZambia
andTanzania(Vollesen2010,2015) which does notextend
toAngola.Theyshare the combination oflanceolateleaves,
eglandularinflorescenceindumentumandprominently3-veined
calyx lobes, bracts and bracteoles. This latter character is
widespread in the perennial species of Justicia sect.Monechma
GroupIsensuKieletal.(2017;seeDiscussionbelow)butis
particularlymarkedinthesetwospecies.However,J. eriniae
clearly differs in the characters listed in the diagnosis, and also
occursinmuchdrierhabitat.
DISCUSSION: A CONSPECTUS OF
JUSTICIA SECT. MONECHMA ‘GROUP I’ IN ANGOLA
Makholela(2008)recorded10speciesofMonechma inAngola.
Of these, M. cleomoides (S.Moore)C.B.Clarke,M. divaricatum
(Nees)C.B.ClarkeandM. salsola (S.Moore)C.B.Clarke are
considered to belong to Monechma GroupIIsensuKieletal.
(2017), based on the sessile axillary cymes with the bracts
largelyundifferentiatedfromtheleaves.Monechma divaricatum
was one of the two species sampled within this clade in the phy-
logenyofKieletal.(2017).TheextantmaterialofM. spissum
C.B.Clarkeisdepauperateandrequiresfurtherinvestigation;it
maybeasmallformofoneoftheotherrecognisedspecies.The
remainingsixspeciesrecordedbyMakholela(2008)arehere
considered to belong to Monechma GroupIsensuKieletal.
(2017).However,severalchangesandadditionstoMakholela’s
listarerequired.First,wefollowVollesen(2010,2015)inrecog-
nisingseveralspecieswithinthe‘Monechma debile (Forssk.)
Nees’complex,ofwhichtwoarerecordedfromAngola:Justicia
bracteata and J. monechmoides S.Moore.Second,Makholela
(2008)recorded M. depauperatum (T.Anderson)C.B.Clarke
(=J. depauperata T.Anderson)asoccurringinAngolabutwe
recognise J. scabrida S.Moore as distinctfrom that species
andonly thelatter occursinAngola (Vollesen2015). Third,
J. subsessilis Oliv.hasrecentlybeenrecordedforthefirsttime
inAngola–seebelow.Finally,J. laeta S.Moore,alsoendemic
toAngola,ishereincludedinthislineageforthefirsttime,and
Character Justicia eriniae Justicia glaucifolia Justicia tricostata
Stemindumentumorientation Patent Antrorse Antrorse
Leaf shape Lanceolate Obovate Lanceolate, narrowly ovate or narrowly
 (oblong-)elliptic,lowerleavescanbe
somewhat obovate or oblanceolate
Leaflength:widthratio 4.5– 6.7:1 2.5 – 2.6:1 3.2– 6 (–8):1
Leafapex Acute Roundedorobtuse Acute,lowerleavessometimesobtuseor
rarely rounded
Inflorescencecharacteristics Laxlyspiciform Laxlyspiciform Denselyspiciformthroughoutoratleastdistally
Bracteolelength 4.5– 6mmlong,±halfaslong 1.8– 2.3mm,±onethirdthe 6.5 –12.5mmlong,slightlyshorterthanor
ascalyx lengthofthecalyx subequalinlengthtocalyx
Calyxlobelength 8– 9mm ±6mm 7.3–11mm
Corollalength 18–18.5mm ±14mm 17–20mm
Capsuleindumentum Puberulous Unknown Glabrous
Seedcharacteristics Black Unknown Mottledgreyandblack
Table 2 Morphological characters separating Justicia eriniae from J. glaucifolia and J. tricostata.
104 Blumea–Volume64/2,2019
a second species allied to J. laeta is also recognised here,
based on Nolde 229fromMalangeprovince.
Therefore,withtheadditionofthenewspeciesJ. cubangen-
sis and J. eriniae, twelve species of Justicia sect.Monechma
GroupIarecurrentlyknownfromAngolaandaprovisionalkey
tothesespeciesis providedheretoaidtheiridentification.A
summary of our current knowledge of each of these species
in Angola is also provided below, including citation of all speci-
mensseen.NoexistingcombinationsinJusticia are available
for three Angolan endemic species: M. glaucifolium S.Moore,
M. rigidum S.Moore and M. virgultorum S.Moore;the new
combinations(or,inthecaseofM. rigidum,anewname)are
thereforeformalised here.Thismaybea temporarysolution
in view of the uncertainty over the future generic delimitation
inthe‘justicioid’lineage, but as we currentlyacceptJusticia
sect.Monechma (seeIntroduction),itissensibletomakethe
requiredcombinationsinJusticia for all the Angolan species.
KEY TO JUSTICIA SECT. MONECHMA GROUP I
IN ANGOLA
1. Annualherbs,inflorescencespikesaxillary,sometimesalso
terminal but those at lower nodes often longest; bracts elliptic
or ovate to broadly so...........................2
1. Perennialherbs,inflorescencespikesorpanicles primarily
terminal, sometimes also in the upper leaf axils; bracts linear,
lanceolate or elliptic .............................3
2. Bract apexsubacute totruncate, markedlyrecurved, with
distinct‘shoulders’;marginwithconspicuouspalelongeglan-
dular hairs contrasting with short indumentum of external
surface .............................1.J. bracteata
2. Bract apex acute and straight or acuminate and slightly
recurved,usuallywithout‘shoulders’,marginsusuallywith-
out markedly longer hairs or if these present then not so
conspicuous and also present on external surface......
..............................8.J. monechmoides
3. Bractsandbracteoleslongerthancalyceswhicharelargely
hidden;inflorescencespikesnot(ornotclearly)secund . 4
3. Bractsandbracteolesshorterthan orbracteolessubequal
in length to calyces which are clearly visible; inflorescence
spikes often markedly secund.....................5
4. Bracts,bracteolesandcalyceswithabroadandconspicuous 
pale hyaline margin, this often drying yellowish; leaves nar-
rowlyelliptictolanceolate,7–14.5cmlong,apexmarkedly
acute;calyxlobes6–7(–9)mmlong ......10.J. scabrida
4. Bracts,bracteoles andcalyces lackingorwith onlya very
narrowhyalinemargin;leavesovateorelliptic,3–7cmlong,
apexobtuseormorerarelyacute;calyxlobes11–19(–21)
mm long .........................11.J. subsessilis
5. Indumentumof bracts, bracteolesand calyceseglandular
only or largely glabrous, glandular hairs absent.......6
5. Indumentumof bracts,bracteoles andcalyceswith mixed
glandular and eglandular hairs ....................9
6. Leaves large, 95 –115 by 38 – 46 mm, elliptic or oblong-
elliptic, apex acuminate................... 5.J. laeta
6. Leavessmaller,33 – 68by 2–14 mm,linear,lanceolate or
obovate, apex acute, obtuse or rounded .............7
7. Bracts,bracteoles andcalyx lobesglabrousexcept forin-
conspicuous ascending hairs along margins; leaves linear
tonarrowlyoblong,upto8mmwide,length:widthratioof
longestleaves3–16:1;capsuleessentiallyglabrous...
....................................7.J. lolioides
7. Bracts, bracteoles and calyx lobes pubescent or puberulous;
leaves obovate or if lanceolate and length : width ratio over
3:1thencapsulepuberulous.....................8
8. Leavesobovatewithapexroundedorobtuse;length:width
ratio2.5–2.6:1;indumentumofstems,inflorescencera-
chisandcalycesantrorse;calyxlobes±6mmlong,corolla
±14mmlong...................... 4.J. glaucifolia
8. Leaveslanceolatewithapexacute,length:widthratio4.5–
6.7:1; indumentum of stems, inflorescence rachis and
calycesspreading/patent;calyxlobes±8mmlong;corolla
±18.5mmlong.......................3.J. eriniae
9. Leaveslinear-lanceolate,2.3 –3.2mmwide,length:width
ratio 10–16 : 1, primary venation of 5 closely parallel veins
...............................2.J. cubangensis
9. Leavesovate,ellipticornarrowlyoblong-elliptic,≥15mm
wide, length : width ratio < 5 : 1, primary venation either
pinnate or campylodromous, the latter with strongly ascend-
ing and converging secondary veins and with prominent
scalariform tertiary veins.......................10
10. Leavesnarrowly oblong-elliptic,upto13 by4.2cm with
base long cuneate-attenuate, primary venation pinnate;
corolla18–20mmlong............6.J. sp.aff. laeta
10. Leaves ovate, elliptic, or elliptic-lanceolate, up to 5 by
2.3cmwith base cuneate torounded,primary venation
campylodromous;corolla11–13mmlong .......... 11
11. Leaves elliptic or elliptic-lanceolate; calyces clearly
longer than bracteoles, lobes oblong-spathulate, markedly
3-veined,theveinsprominentandpurpleinfreshmaterial;
inflorescencespikesatfirstcrowdedbutproximalflowers
becoming more widely spaced...........9.J. moorei
11. Leaves ovate; calyces only slightly longer than bracteoles,
lobeslinear-lanceolate, only inconspicuously 3-veined,
only the lobe apices coloured purple; inflorescences spikes
crowded throughout .............. 12.J. virgultorum
1. Justicia bracteata (Hochst.)Zarb
Justicia bracteata (Hochst.) Zarb (1879) 32.—Monechma bracteatum
Hochst.(1841)375.—Type:Kotschy 261 (holoTUB[notseen];isoGOET*
[GOET005559], GZU [not seen], HBG* [HBG502284, HBG502285], K
[K000378692], LD* [LD1226235], M* [M0109807], MPU [not seen], S*
[S09-6163],STU* [STU000468], US*[US01049972]),Sudan, Kordofan,
Tejara,19Nov.1839.
Monechma debile sensuauct.,non (Forssk.)Nees:Meyer(1968)45p.p.;
Makholela(2008)23p.p.;Munday(1995)59p.p.
 Distribution&Ecology—Thisisawidespreadandcommon
species,recordedthroughouteasternandsouthernAfrica(An-
gola,Botswana,D.R.Congo,Eritrea,Ethiopia,Kenya,Malawi,
Mozambique,Namibia,Somalia,SouthAfrica,Sudan,Tanza-
nia,Uganda, Zambia, Zimbabwe), southernArabia(Oman,
Yemen)andIndia.InAngola,itisrestrictedtothesouth,having
beenrecordedinCuandoCubangoandHuílaprovinces.Itis
an annual herb, found in a variety of open to lightly shaded
habitats including woodland, bushland and grassland, often in
disturbedareaswhereitcanbeaweed.
Angolan specimens seen.Baum 775(BM,K),CuandoCubango,zw.Kutue
undSobiamCuito[Kuito],fl.&fr.,10Mar.1900;Barbosa & Gouveia 10692
p.p. (BM),Huíla,Gambos,Chimbolelo,fl.&fr.,24Aug.1963;Harris & Murray-
Hudson 860 (K),CuandoCubango,CuitoR.,betweenRitoandNankova,
fl.&fr.,19Apr.2013.
 Notes—ThisspecieshascommonlybeennamedMonech-
ma debile inthepast.However,M. debile s.str. (=Justicia debilis
(Forssk.)Vahl)isaperennial(rarelyannual)herbofNortheast
AfricaandArabia(Vollesen2010).
For a full synonymy and bibliography for this species, see
Vollesen(2010,2015).
2. Justicia cubangensis I.Darbysh.&Goyder
SeeNewSpeciesaccountsabove.
105
I.Darbyshire&D.J.Goyder:Justicia section Monechma in Angola
3. Justicia eriniae I.Darbysh.
SeeNewSpeciesaccountsabove.
4. Justicia glaucifolia (S.Moore)I.Darbysh.&Goyder,comb.
nov.
Monechma glaucifolium S.MooreinJ.Bot.49(1911)310;S.Moore(1930)
138. — Type:Gossweiler s.n.(holoBM[BM000923665]),Angola,without
preciselocalityordate.
 Distribution&Ecology—Justicia glaucifolia is known only
from the type specimen, for which the collection data were
lost.Theexactdistributionandtheecologyofthisspeciesare
therefore unknown, but it is currently considered to be endemic
toAngola.
5. Justicia laeta S.Moore
Justicia laeta S.Moore(1880) 311;Hiern(1900)820. — Type:Welwitsch
5108 (lectoBM*[BM000839214],chosenhere;isolectoK[K000419259]),
Angola,PungoAndongo,interCondoetQuisonde,fl.,Mar.1857.
 Distribution&Ecology—Justicia laeta is known only from
the two syntype gatherings from Malange province, collected
frombushyhillslopesandamongstrocks(Hiern1900).
Additional Angolan specimen seen.Welwitsch 5081 (BM), PungoAn-
dongo,FontedeCasamba,fl.&imm.fr.,May1857.
 Note—Moore(1880) suggested that his J. laeta is allied
to J. petiolaris (E.Mey.exNees)T.Anderson,butdidnotela-
borateonthisaffinity.Justicia petiolaris belongswithin sect.
Tyloglossa (Hochst.)Lindau(Vollesen2015,Kieletal.2017)
and is morphologically very different to J. laeta. TheBMsyntype
Welwitsch 5081 includes an immature capsule with poorly de-
velopedseedsthatappeartobesmoothasinsect.Monechma.
This,togetherwiththeprominently3-veinedcalyxlobes,bracts
and bracteoles, and the fact that some of the spikes of the
inflorescence are secund, indicates that this species belongs
insect.Monechma GroupI.Seealsonotetospecies6below.
6. Justicia sp.aff.laeta S.Moore
 Distribution&Ecology—Thistaxon isknownonlyfroma
single collection from Malange province, where it was recorded
fromrainforest.
Angolan specimen seen.Nolde 229(BM2sheets),hochlandvonQuela,
fl.&fr.,withoutdate.
 Notes—Thespecimen cited has previouslybeennamed
J. laeta and is certainly closely allied to that species, sharing
large leaves with prominent pinnate veins beneath, compound
paniculateinflorescences andsimilarly sizedcorollas. How-
ever, Nolde 229 differs in having narrower, linear-lanceolate
bracts,bracteolesandcalyxlobes,thelatter c. 1.2 –1.5 mm
wide, and in the inflorescence having a mixed glandular and
eglandular indumentum, the glandular hairs being longer and
conspicuous, while in J. laetas.str. the bracts and bracteoles
are lanceolate-acuminate, the calyx lobes are 1.8 –2.5 mm
wideandtheinflorescencehasaneglandularindumentumonly.
Thespecimenwascollectedc.100–170kmtotheENEofthe
known localities for J. laeta and apparently in a wetter, forest
habitat. More material is desirable from this region of Angola
to determine whether there are two species or one variable
speciesinvolved.
One sheet of Nolde 229 includes a mature fruit which appears
to be four-seeded but with two seeds aborting early; the mature
seeds are unfortunately missing from this specimen but the
immature seeds are smooth as in J. laeta and other members
of Justiciasect.Monechma.Thiscapsuletypeissimilartothat
of J. tetrasperma,thespeciesthatHedrén(1990)considered
to be intermediate between Justicia and Monechma (see
Introduction).
7. Justicia lolioides S.Moore
Justicia lolioides S.Moore (1880)310.—Monechma lolioides (S.Moore)
C.B.Clarke(1900) 218; Hiern (1900)822. — Type: Welwitsch 5178
(lecto BM [BM000923676], chosen here; isolecto C* [C10000041], K
[K000378722],LD*[LD1569502],LISU*[LISU223483]),Angola,Malange,
Dist.PungoAndongo,deMatadeMutollo,fl.&fr.,Jan.1857.
1. Leaves linear, length:width ratio 18– 24:1...........
.................................. a.var. lolioides
1. Leaves oblong, length:width ratio 3 –6.5:1 ...........
...................................b.var.latifolia
a.var.lolioides
 Distribution& Ecology—Justicia lolioides var.lolioides is
endemictoAngolawhereitisrecordedfromMalangeprovince.
It is only known from the two original syntypes; Welwitsch
recordeditfrom‘moist,hotwoodedpartsofMatadeMutollo’
(Hiern1900:822).
Additional Angolan specimen seen. Welwitsch 5090(originalsyntypes
BM,K,LISU*),PungoAndongo,fr.,withoutdate.
b.var.latifolia S.Moore
Justicia lolioides var.latifolia S.Moore(1880)310.—Type:Welwitsch 5099
(lectoBM [BM000923675] chosen here; isolecto C* [C10000039], K
[K000378723],LD*[LD1575500]),Angola,Malange,Dist.PungoAndongo,
interQuisondeetCondo,fl.&fr.,Mar.1857.
 Distribution& Ecology—Justicia lolioides var.latifolia is
endemic to Angola and is known only from the type specimen
from Malange province, where it was recorded as growing in
dense masses in somewhat dry wooded pastures with sparse
herbage(Hiern1900).
Notes — Clarke(1900:218)recorded this species as oc-
curringin‘PungoAndongoandHuila’butthisisinerrorasthe
syntypesofvar.lolioides (Welwitsch 5090, Welwitsch 5178)
andthetypeofvar.latifolia(Welwitsch 5099)areallfromthe
Pungo Andongo region, and there are no further collections
known.ThefoliageisrathervariableinJ. lolioides.Theleaves
ofvar.lolioides are linear as in J. cubangensis, but those of
var.latifolia areoblong,upto8mmwidewithalength:width
ratioof±3.5–6:1andwithpinnatevenation.
8. Justicia monechmoides S.Moore
Justicia monechmoides S.Moore(1880)311.—Monechma monechmoides
(S.Moore)Hutch.(1946)524.—Type:Welwitsch 5140 (lectoBM,chosen
byVollesen2015:214;isolectoK,LISU*[LISU223491,LISU223493],P*
[P00434925]),Angola,Luanda,ImbondeirodosLobos,fl.&fr.,Mar.1858
[datefideHiern1900].
Monechma welwitschii C.B.Clarke(1900)216,nom.illegit;Hiern(1900)822;
Moore(1930)137;Benoist(1950)29.—Syntypes:Welwitsch 5065 (BM,
K),Welwitsch 5123 (BM),Welwitsch 5140(BM,K,LISU*),Welwitsch 5184
(K),Angola,Loanda,fl.&fr.,Mar.1858[datefideHiern1900]–seenote.
Monechma tettense C.B.Clarke (1900) 216.—Type:Kirk s.n. (lecto K
[K000378730],chosenbyVollesen2015:214),LowerZambesi,opposite
Tete,KaimbaIs.,1860.
Monechma debile sensuauct.,non (Forssk.)Nees:Meyer(1968)45p.p.;
Munday(1995)59p.p.;Makholela(2008)23p.p.
 Distribution&Ecology—Justicia monechmoides is wide-
spread in southern Africa, occurring in Angola, Botswana,
Malawi, Mozambique, Namibia, South Africa, Zambia and
Zimbabwe.InAngola, it is recordedfromCuandoCubango,
Cunene,Huíla,LuandaandNamibeprovinces.Itisanannual
106 Blumea–Volume64/2,2019
herb of a variety of woodland, bushland and grassland habitats,
oftenfavouringdisturbedareaswhereitcanbeaweed.
Additional Angolan specimens seen.Welwitsch 5065(BM,K),5123 (BM),
5184 (K),Luanda[Loanda],ImbondeirodosLobos,fl.&fr.,Mar.1858[precise
localityanddatefideHiern1900];Gossweiler 306(BM,K),306B (BM),307
(BM,K),Luanda[Loanda],fl.&fr.,1903;Pearson 2684 (K),Huíla,between
ChibiaandQuihila,fl.&fr.,13May1909;Pearson 2221 (K),betweenQui-
hila-Jambos, 3–18miles,fl.&fr.,14May1909;Exell & Mendonça 2403
(BM),2413(BM),Namibe,Moçâmedes,Tampa,fl.&fr.,1June1937;Exell
& Mendonça 2958(BM), Huíla, near Leba, fl.&fr.,20June1937;Goss-
weiler 14268(BM),Luanda,Catete,fr.,July1949;Pritchard 357(BM),Huíla,
Tchivinguiro,fl.&fr.,3Aug.1954;Barbosa & Gouveia 10692p.p. (BM),Huíla,
Gambos,Chimbolelo, fl. & fr., 24Aug.1963; Borges 227 (LISC*), Huíla,
Lubango,aos600mdapicadaparaamissãodoMunhino,fl.&fr.,11May
1971; Santos 2910 (LISC*),Cunene,Cuvelai,fl.&fr.,21July1971;Harris
& Murray-Hudson 828 (K),Cuando Cubango, Rápidos M’Pupo,CuitoR.,
betweenDingoandLumeta,fl.&fr.,17Apr.2013;Harris & Murray-Hudson
1028 (K),CuandoCubango,20kmSEofSiangaonSWedgeofR.Cuando
floodplainnearBenorio,fl.&fr.,28Apr.2013.
 Notes—Thisspeciesissometimestreatedasconspecific
with J. bracteata (underthenameMonechma debile – see note
under J. bracteata above) and the two are certainly close but
areusuallyreadilyseparatedbythecharacterslistedinthekey.
The five Welwitsch syntypes listed for Clarke’s (1900) Mo-
nechma welwitschiiarethesameasthoseofMoore’s(1880)
Justicia monechmoides;Vollesen(2015)selectedWelwitsch
5140 as the lectotype of J. monechmoides and noted under
M. welwitschii (p.214):“type:asforJusticia monechmoides”,
buthedidnotformallylectotypifythisname.
9. Justicia moorei I.Darbysh. & Goyder, nom. nov.—Fig.
2d–e
Monechma rigidum S.MooreinJ.Bot.49(1911)310;S.Moore(1930)138.
—Type:Gossweiler 3355(lectoBM[BM00092362],chosenhere;isolecto
K[K001009703]),Angola,Menongue[Munongue],fl.,Apr.1906.
Etymology.Theepithet rigida is already occupied in Justicia(J. rigida
Balf.f.)andsoanewnameisrequiredforthisAngolanspecies.Theepithet
‘moorei’, honours Spencer leMarchantMoore(1850–1931)whoworked
extensively on the Acanthaceae of Africa amongst many other plant groups,
and who originally described this species as Monechma rigidum.
 Distribution&Ecology–Justicia moorei is endemic to Angola
whereitisrecordedfromBiéandCuandoCubangoprovinces.
Itoccursindrysunnysitesongravel-claysoils,onshortgrassy
slopes and on raised ground above marshes; Goyder 8210 was
collectedfrom1648melevation.
Additional Angolan specimens seen. Gossweiler 2535(BM[BM000923663],
K[K000378713],originalsyntypes),CuandoCubango,FortePrincesaAme-
lia,Cubango,fl.,27Jan.1907;Mendes 2425 (LUBA*),CuandoCubango,
Menongue[VilaSerpaPinto],fl.,9Feb.1960;Goyder 8210(INBAC,K,LUBA,
PRE),Bié,RíoCutato52kmWofKuito,Cubangodrainage,fl.,11Feb.2016.
 Note—ThereisalsoanunnumberedGossweilerspecimen
at K which appears to be a further duplicate of Gossweiler 3355.
10. Justicia scabrida S.Moore
Justicia scabrida S.Moore(1880)310. — Monechma scabridum (S.Moore)
C.B.Clarke(1900) 217; Hiern (1900) 822;S.Moore(1930)137; Benoist
(1950)29.—Type:Welwitsch 5092(lectoBM[BM000923677],chosen
byVollesen2015:222;isolectoK[K000378724], LISU* [LISU223487]),
Angola,PungoAndongo,fl.,11Mar.1857.
Justicia marginata Lindau(1894)73.—Type:Buchner 33 (holoB†;isoK
[K000378726,K000378727]),Angola,Malange[Malansche],fl.&fr.,Mar.
1879.
Monechma depauperatum sensu Makholela(2008) 23, non (T.Anderson)
C.B.Clarke.
 Distribution&Ecology—Justicia scabrida is recorded from
D.R.Congo,ZambiaandAngola;inAngolaitisfoundinLunda
SulandMalangeprovinces.Itisfoundinfire-pronewoodland
and bushland including miombo woodland and in open, tall
grasslandaswellasinfallowfields.
Additional Angolan specimens seen.Welwitsch 5085 (BM, K, LISU*,
originalsyntype),Dist.PungoAndongo,propeCondo,NbilleetBumba,fl.,
Mar.1857;Gossweiler 1084(BM,K),Loanda,GamanhiangonearMalange,
fl.&fr.,5June1903;Exell & Mendonça 430(BM),LundaSul,Xa-Sengue,
fl.,8Apr.1937;Rocha 123 (LUBA*),Malange,Cambo,Sunginge,fl.,28Mar.
1953.
 Note—Vernacularnameandlocaluses:Moka(Chiokwe),
rubbedonthroat(Exell & Mendonça 430).
11. Justicia subsessilis Oliv.
Justicia subsessilis Oliv.(inGrantetal.1875)129,t.129b.—Monechma
subsessile (Oliv.)C.B.Clarke (1900) 216.—Type: Speke & Grant 213
(holoK[K000378735]),Tanzania,BukobaProv.,Karagwe,fl.,2Dec.1861.
Justicia simplicispica C.B.Clarke(1900)188.—Type:Scott Elliot s.n.(holo
K[K000378734]),EastAfrica,withoutlocalityordate.
Monechma nemoralis S.Moore (1909) 296.—Type:Kässner 3010 (holo
BM[BM000923664];isoK[K000379152]),D.R.Congo,NiembeR.,27
May1908.
 Distribution&Ecology—Justicia subsessilis is widespread
across central and southern tropical Africa, largely associ-
atedwith theZambesian regionalcentre ofendemism. It is
recordedinBurundi,D.R.Congo,Kenya,Rwanda,Tanzania,
Uganda,Zambia andZimbabwe.InAngola,itis knownfrom
CuandoCubangoProvince.Itoccursinfire-pronewoodland,
bushland and short grassland; the single Angolan record was
from1234melevation.
Angolan specimen seen. Barker, Bester & Janks 89(INBAC,K,LUBA,
PRE),CuandoCubango,LongaRiver,55kmSWofCuitoCuanavale[Cuito
drainage],fr.,7June2015.
 Note—ThisspecieswascollectedforthefirsttimeinAn-
gola in 2015 during a survey of the more southerly catchment
of the Cubango and Cuito rivers in Angola by a team from
GrahamstownandPretoria in SouthAfrica, again as partof
theOkavangoWildernessProject.
12. Justicia virgultorum (S.Moore)I.Darbysh.&Goyder,
comb. nov. —Fig.2f
Monechma virgultorum S.MooreinJ.Bot.49(1911)311;S.Moore(1930)138.
—Type:Gossweiler 3679 (holo BM [BM000923667]), Angola, Cuando
Cubango,Cassuango,Cuiriri,fl.,26March1906.
Monechma carrissoi Benoist(1950)30,syn.nov.—Type:Carrisso & Sousa
113 (holoCOI[notseen],?†];isoBM[BM001252126]),Angola,Benguela,
Huambo[NovaLisboa],fl.,18May1937.
Distribution & Ecology—Justicia virgultorum is endemic
toAngola,whereitisrecordedfromBié,Huambo,Huílaand
probably(fideBarbosa & Correia 9051)Benguelaprovinces.It
isrecordedfromdryshortgrasslandandshrubland(‘anharas’)
and from open miombo woodland of Julbernardia and Brachy-
stegia.
Additional Angolan specimens seen. Angola, Hundt 907(BM,2sheets),
Benguela,zwischenGandaundCaconda,FacendaXangorolo,fl.,Apr.1934;
Barbosa & Correia 9051(BM,K),Huíla,Capelongo[VilaArturdePaiva],a
26kmdesta, emdirecção aoDongo,fl.,10Apr.1960;Bamps, Raimundo
& Matos 4007 (LISC*),Huambo [Nova Lisboa] - Caála,fl., 6 Mar.1973;
Maiato FM 897(139013)(K,LUBA),Bié,Cussequestudyarea,Chitembo,
fl.,19Mar.2013;Goyder 8471(INBAC,K,LUBA,PRE),Bié,nearLinhunga
R.,Cubangodrainage,fl.,14Mar.2016; Frisby & Maiato 4151(K, LUBA,
PRE),Huambo,CubangoR.rapids75kmESEofHuamboonCatchiungo
–Kuvango(Capelongo)road,inbud,9May2017;Frisby & Maiato 4183(K,
LUBA,PRE),Huambo,nearCubangoRiverrapids75kmESEofHuambo
onCatchiungo–Kuvango(Capelongo)road,fr.,10May2017.
 Notes—ThisspeciesisalliedtoJ. moorei (seeabove)from
Angola,andP.G.MeyernotedontheBMsheetsthattheyare
scarcelydistinct. However,the current studyfinds thatthey
107
I.Darbyshire&D.J.Goyder:Justicia section Monechma in Angola
clearlydifferinthecharacterslistedinthekey;seealsoFig.
2d–fforcomparisonofthesespecies.ThetypeofMonechma
carrissoi is clearly a specimen of J. virgultorum but it has shorter
calyxlobes(7mmlong)andbracts(c.6mm)thantypical;this
ismostprobablyanimmatureordepauperatespecimen.
TheKsheetofBarbosa & Correia 9051isnumbered‘9051a’
butthecollectingdetailsarethesameasontheBMsheet.
AcknowledgementsWethank Erin Trippof the University ofColorado
for kindly sending a duplicate of the new species Justicia eriniae to K and
forproviding additionalfloralmeasurementsfrom theCOLOisotype. D.J.
GoyderismostgratefultothedirectorsofNationalGeographicOkavango
WildernessProject,WildBirdTrust,SouthAfricaforfacilitatingfieldwork,and
toTheHaloTrustforadviceandlogisticalsupport.WethankJulietWilliamson
fortheexcellentillustrations.Finally,wethanktwoanonymousreviewersfor
theirconstructivecommentsonadraftofthismanuscript.
REFERENCES
AndersonT.1863.AnenumerationofthespeciesofAcanthaceaefromthe
continentofAfricaandtheadjacentislands.JournaloftheProceedingsof
theLinneanSociety,Botany7:13–54.
BadenC,BalkwillK,GetliffeNorrisFM,etal.1995.Acanthaceae.Fascicle
1:Justiciinae (Metarungia, Siphonoglossa, Rhinacanthus, Duvernoia,
Justicia,Monechma,AdhatodaandAnisotes).In:LeistnerOA(ed),Flora
ofSouthernAfrica,Vol.30,Part3.NationalBotanicalInstitute,Pretoria.
BenoistR.1950.QuelquesAcanthacéesdescoloniesPortugaisesAfricanes.
BoletimdaSociedadeBroterianaSér.2,24:5– 39.
ClarkeCB.1900.Monechma.In:Thiselton-DyerWT(ed),FloraofTropical
Africa.Vol.5:212– 220.Reeve&Co.,London.
DarbyshireI. 2015. ThegenusHypoestes(Acanthaceae)inAngola.Kew
Bulletin70:44(10pages).
DarbyshireI,GoyderD,CrawfordF,etal.2014.Updatetoreportontherapid
botanicalsurveyoftheLagoaCarumboRegion,LundaNorteProv.,Angola
fortheAngolanMinistryoftheEnvironment,followingfurtherfieldstudies
in2013,incl.Appendix2:checklisttothefloweringplants,gymnosperms
and pteridophytes of Lunda Norte Prov.,Angola. In: Huntley BJ (ed),
Biodiversity rapid assessment of the Lagoa Carumbo area, Lunda Norte,
Angola.Unpubl.Report,Annex3:59– 98.
DarbyshireI,KordofaniM,Farag I,et al.2015a.TheplantsofSudanand
SouthSudan,anannotatedchecklist.RoyalBotanicGardens,Kew.
DarbyshireI,NanyeniL,ChaseFM,etal.2018.AsynopsisofRhinacanthus
(Acanthaceae)inAngolaandNamibia.KewBulletin73:21(12pages).
DarbyshireI,TrippEA,ChaseFM.2019.AtaxonomicrevisionofAcantha-
ceaetribe BarlerieaeinAngolaand Namibia. Part1.Kew Bulletin74:5
(85pages).
DarbyshireI,VollesenK,EnsermuKelbessa.2010.Acanthaceae(Part2).In:
BeentjeH(ed),FloraofTropicalEastAfrica.RoyalBotanicGardens,Kew.
DarbyshireI,VollesenK,EnsermuKelbessa.2015b.Acanthaceae(Part2).
In:TimberlakeJR,MartinsES(eds),FloraZambesiaca.Vol.8(6).Royal
BotanicGardens,Kew.
EnsermuKelbessa.2006.Acanthaceae.In:HedbergI,EnsermuKelbessa,
EdwardsS,etal.(eds),FloraofEthiopia&Eritrea.Vol.5:345–495.The
NationalHerbarium,AddisAbabaUniversity&TheDepartmentofSyste-
maticBotany,UppsalaUniversity.
FigueiredoE,SmithGF.2008.PlantsofAngola/PlantasdeAngola.Strelitzia
22.SouthAfricanNationalBiodiversityInstitute,Pretoria.
FischerE.1989.ContributionstothefloraofCentralAfricaII.Crepidorho-
palon,anewgenuswithintherelationshipofCraterostigma,Toreniaand
Lindernia(Scrophulariaceae) with two new or noteworthy species from
Centraland South Central Africa (Zaire, Zambia). Feddes Repertorium
100:439– 450.
Fischer E. 1992. Systematik der afrikanischen Lindernieae (Scrophula-
riaceae).Tropischeund Subtropische Pflanzenwelt 81. Akademie der
WissenschaftenundderLiteratur,Mainz.FranzSteinerVerlag,Stuttgart.
GhazanfarSA.2008.Scrophulariaceae.In:FigueiredoE,SmithGF(eds),
PlantsofAngola/PlantasdeAngola.Strelitzia22:158 –159.SouthAfrican
NationalBiodiversityInstitute,Pretoria.
GonçalvesFMP,GoyderDJ.2016.Abriefbotanicalsurveyinto Kumbira
forest,anisolatedpatchofGuineo-Congolianbiome.Phytokeys65:1–14.
GoyderDJ,GonçalvesFMP.2019.TheFloraofAngola:collectors,richness
andendemism.In:HuntleyBJ,RussoV,LagesF,etal.(eds),Biodiversity
ofAngola.Scienceandconservation:Amodernsynthesis:79–96.Springer,
Cham,Switzerland.
GrantJA,OliverD,BakerJG,etal.1875.ThebotanyoftheSpekeandGrant
expedition, an enumeration of the plants collected during the journey of
thelateCaptainJ.H.SpekeandCaptain(nowLieut.-Col.)J.A.Grantfrom
ZanzibartoEgypt.TransactionsofLinneanSocietyofLondon29:1–129.
HedrénM.1989.Justicia sect.Harnieria(Acanthaceae)inTropicalAfrica.
SymbolaeBotanicae Upsaliensis 29: 1.Acta Universitatis Upsaliensis,
Uppsala.
HedrénM.1990.Justiciatetraspermasp.nov.:AlinkingspeciesbetweenJus-
ticia and Monechma (Acanthaceae).NordicJournalofBotany 10:149–153.
HiernWPH. 1900. Catalogue of the African plants collected byFriedrich
Welwitschin1853 –1861.Dicotyledons,partIVLentibulariaceaetoCerato-
phylleae.BritishMuseum(NaturalHistory),London.
HochstetterCHF.1841. Plantarum Nubicanumnova genera proponit et
describit.Flora24:369– 377.
HutchinsonJ.1946.AbotanistinSouthernAfrica.GawthornLtd.,London.
IUCN2012. IUCN Red List categories and criteria. Version3.1. Second
Edition.IUCNSpeciesSurvivalCommission,Gland,Switzerland&Cam-
bridge,UnitedKingdom.
KielCA,DanielTF,DarbyshireI,etal.2017.Unravelingrelationshipsinthe
morphologicallydiverseandtaxonomicallychallenging‘justicioid’lineage
(Acanthaceae:Justicieae).Taxon 66:645– 674.
KimpouniV.1994.NouvellesouintéressantesespècesdugenreEriocau-
lon(Eriocaulaceae)enAfrique.FragmentaFloristicaetGeobotanica39:
319– 359.
LindauG.1894.AcanthaceaeafricanaeII.BotanischeJahrbücherfürSys-
tematik,PflanzengeschichteundPflanzengeographie 20:1–76.
MakholelaT.2008.Acanthaceae.In:FigueiredoE,SmithGF(eds),Plants
ofAngola/PlantasdeAngola.Strelitzia22:21– 25.SouthAfricanNational
BiodiversityInstitute,Pretoria.
ManktelowM.1996.Phaulopsis (Acanthaceae)–amonograph.Symbolae
BotanicaeUpsaliensis31:2.ActaUniversitatisUpsaliensis,Uppsala.
ManningJC,GoldblattP.2014.NomenclaturaladjustmentsinAfricanplants
1.1.SynopsisofthegenusJusticiaL.(Acanthaceae)insouthernAfrica,
with eight new combinations for species of Adhatoda Miller and Monechma
Hochst.Bothalia 44:42– 47.
MeyerPG.1968.130.Acanthaceae.In:MerxmüllerH(ed),Prodromuseiner
FloravonSüdwestafrika.Cramer,Germany.
MooreSleM.1880.EnumeratioAcanthacearum Herbarii Welwitschiani
Angolensis.JournalofBotany18:193 –199,225–233,265–270,307–314,
340– 342,362 – 366.
MooreSleM.1909.AlabastraDiversa.PartXVIII.3.NewTropicalAfrican
Acanthaceae.JournalofBotany 47:293– 296.
MooreS le M. 1911.Alabastra diversa. PartXXI.AcanthaceaeAfricanae
Novae.JournalofBotany49:293– 298,305 – 313.
MooreSleM.1930.MrJohn Gossweiler’s plantsfromAngola andPortu-
gueseCongo.Acanthaceae.JournalofBotany 68,Suppl.II:126–139.
MundayJ.1995.Monechma.In:LeistnerOA(ed),FloraofSouthernAfrica,
Vol.30,Part3:47–61.NationalBotanicalInstitute,Pretoria.
PhillipsSM.2010.Eriocaulaceae.In:TimberlakeJR,MartinsES(eds),Flora
Zambesiaca13(4):33– 86.RoyalBotanicGardens,Kew.
RomeirasMM, Figueira R,DuarteMC,et al. 2014. Documentingbiogeo-
graphical patterns of African timber species using herbarium records: a
conservationperspectivebasedonnativetreesfromAngola.PLoSONE
9(7):e103403. doi: https://doi.org/10.1371/journal.pone.0103403. [Last
accessed25Jan.2018.]
ThiersB.[continuouslyupdated].IndexHerbariorum: aglobal directoryof
publicherbariaandassociatedstaff.NewYorkBotanicalGarden’sVirtual
Herbarium.http://sweetgum.nybg.org/science/ih/.[Lastaccessed25Jan.
2018.]
TrippEA, TsaiY-HE, Zhuang Y, et al. 2017. RADseq dataset with 90%
missing data fully resolves recent radiation of Petalidium(Acanthaceae)
intheultra-ariddesertsofNamibia.Ecology&Evolution7: 7920–7936.
VanWykAE,SmithGF.2001.Regions of floristic endemisminSouthern
Africa.Areviewwithemphasisonsucculents.UmdausPress, Hatfield,
SouthAfrica.
VollesenK. 2000.Blepharis (Acanthaceae).Ataxonomicrevision. Royal
BotanicGardens,Kew.
VollesenK.2010.Justicia.In:BeentjeH(ed),FloraofTropicalEastAfrica:
495–601.RoyalBotanicGardens,Kew.
VollesenK.2015.Justicia.In:TimberlakeJR,MartinsES(eds),FloraZambe-
siaca.Vol.8(6):162–224.RoyalBotanicGardens,Kew.
ZarbJH.1879.Rapportsurlesspécimensbotaniquescolligéspendantles
expéditionsÉgyptiennes auKardofanet auDarfuren 1875 et1876par
leDr.Pfund, naturaliste. Imprimerie de l’État-Major Général Egyptien,
Cairo.
... Although recent collections and descriptions of new species are adding more and more information (e.g. Hind and Goyder 2014;Darbyshire and Goyder 2019), knowledge on floristic diversity in Angola is still far from complete. The "Angolan Protected Area Expansion Strategy -APAES" (Huntley 2010) highlighted 11 areas of particular biological significance. ...
... The checklist of Figueiredo and Smith (2008) was used as the baseline for assessing Angolan species records. Species missing there but subsequently reported in later publications (Swanepoel 2009;Bergh and Nordenstam 2010;De Sousa et al. 2010;Catarino et al. 2013;Abreu et al. 2014;Hind and Goyder 2014;Paton 2014;Cheek et al 2015;Darbyshire 2015;Abrahamczyk et al. 2016;Darbyshire et al 2018;Harris and Wortley 2018a, 2018b, 2018cDarbyshire and Goyder 2019;Swanepoel 2019;Tripp and Darbyshire 2020;Lautenschläger et al. 2020b;Darbyshire et al. 2021;Smith and Lautenschläger 2021;Swanepoel et al. 2021, Figueiredo andSmith 2022) were excluded. ...
Sixty-seven species newly recorded for the flora of Angola: recent findings from vegetation surveys and herbarium collections
Article
Full-text available
  • Nov 2022
Angola is a tropical country with many biogeographical units and, therefore, has a high floristic diversity. Although an increasing number of floristic studies has been carried out in Angola in recent years, the country is still considered to be underinvestigated as many species being collected were previously unknown there. Several scientific groups working in different parts of Angola contributed to this paper their data from biodiversity assessments. With this we can add 67 species newly recorded for Angola, including two new generic records and five alien species, to the almost 7,300 vascular plant taxa known so far for Angola. Most of the new records for Angola are also present in different neighbouring countries, but they are little known, and their IUCN threat status has not been assessed yet. However, ongoing fieldwork and exploration are needed to complete the floristic knowledge of the understudied country.
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... cleomoides (S.Moore) C.B.Clarke and relatives) primarily from the succulent biome of southern Africa (Darbyshire et al. 2020). This tropical savanna group was not sampled in the earlier study of Kiel et al. (2017) but based on morphological extrapolation, both that study and Darbyshire & Goyder (2019) later surmised that the tropical savanna group was allied to species of the M. debile (Forssk.) Nees complex, which are mostly annual, ruderal species with spiciform inflorescences and highly modified bracts. ...
... Species delimitation follows a combination of Hedrén (2006aHedrén ( , 2006b, Munday (1995), Vollesen (2010Vollesen ( , 2015 and Darbyshire & Goyder (2019), with some modifications based on the current study. For each accepted species, the geographic range is derived from herbarium collections, online databases (notably Plants of the World Online, POWO 2020) and relevant taxonomic literature, principally those cited above. ...
The demise of Monechma: new combinations and a new classification in the resurrected genera Meiosperma and Pogonospermum (Acanthaceae)
Article
Full-text available
  • Apr 2022
Recent molecular phylogenetic results have demonstrated that Monechma s.l., a group of plants with ecological importance in the savanna and succulent biomes of sub-Saharan Africa, is polyphyletic with two discrete lineages recognisable. In the present work, we recognise Monechma Groups I and II at the generic rank, which can be distinguished by differences in inflorescence characteristics and seed morphology. The nomenclatural implications of these findings are investigated. The lectotype of Monechma , M. bracteatum Hochst., is a part of a small lineage of plants closely allied to Justicia L. sect. Harnieria (Solms) Benth. for which the earliest valid name is found to be Meiosperma Raf. Hence, Monechma is synonymised within Meiosperma , which comprises six accepted species and two undescribed taxa. The majority of species of former Monechma s.l. are resolved within the second lineage for which the only validly published generic name is Pogonospermum Hochst. This resurrected genus comprises 34 accepted species plus two undescribed taxa. Pogonospermum displays considerable morphological variation and is here subdivided into six sections based primarily on differences in plant habit, inflorescence form, calyx, bract and bracteole venation, and seed indumentum. The new combinations and new sections are validated, and seven accepted species names are lectotypified.
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... Angola has a large diversity of ecosystems and a rich and diverse indigenous flora, with about 6 850 native vascular plant species recorded (Figueiredo et al. 2009). However, the number of species of vascular plants may be much higher; in recent years several species have been described from Angola and numerous new country-level records have been published (e.g., Catarino et al. 2012, Abreu et al. 2014, Darbyshire & Goyder 2019, Goyder & Gonçalves 2019. ...
An annotated checklist of the vascular flora of Quiçama National Park, Angola
Article
  • Aug 2022
The Quiçama National Park (PNQ), located in Luanda Province, Angola, is one of the oldest protected areas in the country. Although some work has been done on the flora of the Park, up to now a comprehensive record of the vascular plant species has been lacking. The objective of the present study was to produce an inventory of the vascular plant diversity of PNQ, as well as to record the types of vegetation in which each species occurs, their habits, uses by local communities, common names, and conservation status. A total of 452 species were recorded, from 285 genera and 85 families. Two species are Pteridophytes, 99 are Monocotyledons, and 351 are Dicotyledons. The two largest families of the PNQ flora are the Fabaceae and the Poaceae. The Park hosts 14 species endemic to Angola, and one of them is restricted to this protected area. Of the 128 species with data on conservation status, 109 are considered of least concern, 13 are vulnerable, and six are endangered. Regarding useful plants, 237 species from 67 families were recorded. Of these, 196 are Dicotyledons and 41 are Monocotyledons. Most of them are used for medicinal purposes, the manufacturing of artefacts, and as sources of food.
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... Additional range-restricted newly described species endemic to Angola are Justicia cubangensis I.Darbysh. & Goyder (Darbyshire & Goyder 2019) and Stomatanthes tundavalaensis D.J.N.Hind (Hind & Goyder 2014). ...
Taxonomic revision of Saxicolella (Podostemaceae), African waterfall plants highly threatened by Hydro-Electric projects
Article
Full-text available
  • May 2022
Species of the genus Saxicolella Engl. (Podostemaceae) are African rheophytes, restricted to rapids and waterfalls as are all members of the family. Previously, Saxicolella sensu lato was shown to be polyphyletic with two separate clades. The name Pohliella Engl. was recently resurrected for one clade that is sister to the American genera Ceratolacis (Tul.) Wedd., Podostemum Michx. and all Old World Podostemoideae. Pohliella has distichous phyllotaxy, bilocular ovaries, filiform roots with paired holdfasts, and rootcaps. The second clade, Saxicolella sensu stricto , including the type of the generic name, has spiral phyllotaxy, unilocular ovaries, ribbon-like or crustose roots that lack both holdfasts and rootcaps. Saxicolella sensu stricto , sampled from the type species, S. nana Engl. of Cameroon, is embedded within and near the base of the major clade of African podostemoids and is sister to all other African genera apart from Inversodicraea R.E.Fr. and Monandriella Engl. Recently reduced to three species in Cameroon and SE Nigeria by the resurrection of Pohliella, Saxicolella sensu stricto is expanded to eight species in this monograph by description of five new taxa. Saxicolella futa Cheek and S. deniseae Cheek are newly described from Guinea, S. ijim Cheek from Cameroon, the informally named S. sp. A from Gabon, and S. angola Cheek from Angola. The known geographic range of the genus is thus expanded c. 2,500 km westwards to Guinea from eastern Nigeria and c.1,500 km southeastwards from near Yaoundé to Cuanza do Sul, Angola. The greatest concentration of species occurs in the Cross-Sanaga interval of western Cameroon and eastern Nigeria, with three species. Cameroon (3 species) followed by Nigeria and Guinea (2 species each) are the countries with highest species diversity. A classification is proposed grouping the species into three subgenera ( Saxicolella, Butumia (G.Taylor) Cheek comb. et stat. nov. and Kinkonia Cheek subgen. nov.) based on root morphology and shoot position and morphology. The discovery, morphology, circumscription, distribution and ecology of Saxicolella is reviewed, an identification key to the species is presented, together with descriptions, synonymy and links to illustrations. All of the species are provisionally assessed as either Endangered or Critically Endangered using the IUCN 2012 Red List Criteria. The major threats, above all, are hydro-electric projects. Saxicolella deniseae may already be globally extinct, and two of the four known locations of S. angola appear lost, S. sp. A of Gabon is threatened at at least one of its three locations, while S. futa is threatened at all three locations. Contamination of watercourses by increased turbidity from silt-load due to anthropic changes and by eutrophication from pollution are also threats for the majority of the species.
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... Durante o desenvolvimento do trabalho foram incluídas na colecção novas espécies para Angola e para a ciência, a maioria colhida durante as expedições botânicas acima mencionadas. No total, registaram-se 38 espécies não constantes da Lista de Plantas de Angola, colhidas em diferentes províncias do país (Gonçalves et al., 2016;Revermann et al., 2017;Goyder et al., 2018) e 15 espécies novas para a ciência (Bergh & Nordenstam, 2010;Hind & Goyder, 2014;Swanepoel, 2012Swanepoel, , 2014Paton, 2014;Goyder et al., 2018;Darbyshire & Goyder, 2019;Tripp & Darbyshire, 2020). A revisão taxonómica de parte da colecção de Acanthaceae (Géneros Barleria, Justicia, Petalidium) conduzida por I. Darbyshire do Herbário de Kew, resultou na descrição de 5 novas espécies , que durante décadas se mantiveram "anónimas" nos armários do Herbário. ...
Caracterização ecológica de um ecossistema florestal do Elundulua
Conference Paper
Full-text available
  • Mar 2022
O trabalho teve como objectivo fazer a caracterização de um Ecossistema Florestal do Elundulua, localizado no Município sede do Huambo, em termo da sua biodiversidade. O estudo incidiu sobre 12 parcelas de 400 m2, cada. Em cada parcela foram classificadas botanicamente e contabilizados os indivíduos presentes. Com bases nestes foram calculados os parâmetros da Estrutura Horizontal. Os resultados obtidos indicam que o ecossistema florestal do Elundulua apresenta uma baixa biodiversidade, na medida em que apesar de apresentar uma grande riqueza, a abundância não está distribuída de forma uniforme. Com efeito, o Ecossistema Florestal do Elundulua apresenta uma grande riqueza, traduzida pelo número total de espécies, (25) e de indivíduos, 746 por hectare. Por outro lado, o ecossistema em questão apresenta uma fraca abundância, isto é, uma má distribuição dos indivíduos pelas diferentes espécies, havendo espécies com muitos indivíduos e outras com pouquíssimos indivíduos, facto que se traduz numa baixa diversidade. Neste ecossistema dominam as seguintes espécies: Brachystegia hocki, com 57 % do total dos indivíduos, Brachystegia sp., com 10%, Sapium ellipticum com 6%. Há espécies como por exemplo, Syzygium guineense, Brachystegia tamarindoides e Diplorhynchus condylocarpn com uma representatividade de 1%.
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... Durante o desenvolvimento do trabalho foram incluídas na colecção novas espécies para Angola e para a ciência, a maioria colhida durante as expedições botânicas acima mencionadas. No total, registaram-se 38 espécies não constantes da Lista de Plantas de Angola, colhidas em diferentes províncias do país (Gonçalves et al., 2016;Revermann et al., 2017;Goyder et al., 2018) e 15 espécies novas para a ciência (Bergh & Nordenstam, 2010;Hind & Goyder, 2014;Swanepoel, 2012Swanepoel, , 2014Paton, 2014;Goyder et al., 2018;Darbyshire & Goyder, 2019;Tripp & Darbyshire, 2020). A revisão taxonómica de parte da colecção de Acanthaceae (Géneros Barleria, Justicia, Petalidium) conduzida por I. Darbyshire do Herbário de Kew, resultou na descrição de 5 novas espécies , que durante décadas se mantiveram "anónimas" nos armários do Herbário. ...
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... Durante o desenvolvimento do trabalho foram incluídas na colecção novas espécies para Angola e para a ciência, a maioria colhida durante as expedições botânicas acima mencionadas. No total, registaram-se 38 espécies não constantes da Lista de Plantas de Angola, colhidas em diferentes províncias do país (Gonçalves et al., 2016;Revermann et al., 2017;Goyder et al., 2018) e 15 espécies novas para a ciência (Bergh & Nordenstam, 2010;Hind & Goyder, 2014;Swanepoel, 2012Swanepoel, , 2014Paton, 2014;Goyder et al., 2018;Darbyshire & Goyder, 2019;Tripp & Darbyshire, 2020). A revisão taxonómica de parte da colecção de Acanthaceae (Géneros Barleria, Justicia, Petalidium) conduzida por I. Darbyshire do Herbário de Kew, resultou na descrição de 5 novas espécies , que durante décadas se mantiveram "anónimas" nos armários do Herbário. ...
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... Generally, the Monechma species are described as well-adapted plants, as they can to survive in hard environments. As reported by Darbyshire and Goyder[6], twelve species are recorded in Angola, with two of them recently added to Monechma. The same authors reported that there are make it traditionally useful for many African populations, especially in rural areas. ...
A review on the health benefits of Monechma ciliatum (Black mahlab): A potential functional food
Article
Full-text available
  • Feb 2022
Background: Monechma ciliatum (MC) (Acanthaceae family) is a herb that grows in tropical and subtropical regions. It grows with small, brownish-black seeds. This is why the Sudanese people call it the “black mahlab”. This plant is primarily used in traditional meals, as well as the production of medical treatments and perfumes. Additionally, the seeds of the Monechma ciliatum plant are rich in fats, proteins, minerals, and other essential nutrients. It also functions as an effective laxative. As a result, this plant is commonly found as a supplement in cereal to enrich nutritional value. This review explores the nutritive value, antioxidants, antimicrobial, medicinal, cosmeceutical, and oxytocic properties of Monechma ciliatum seeds, leaves, and stem. To prepare this review, we researched published scientific literature by using the following keywords: “Monechma”, “Antioxidants”, “Antimicrobial”, nutritional”, “Monechma ciliatum”, “Monechma phenolics”. Our search was carried out in PubMed and Google Scholar databases and included original research from the past 25 years in English only. A total number of 50 original papers, which mentioned the phytochemistry, nutritional, biological, and medicinal properties of Monechma ciliatum were selected and considered for this review. Keywords: Monechma ciliatum, black mahlab, nutritional, antioxidants, Antimicrobial, oxytocic
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... Monechma plants are well adapted to harsh environments. As reported by Darbyshire and Goyder, twelve species are recorded in Angola, two of which have recently been added to Monechma [6]. Although these two species are morphologically similar, especially in flower and fruit morphology, there is some morphological evidence to support their separation. ...
Black mahlab (Monechma ciliatum L.) seeds: processing effects on chemical composition and nutritional value
Article
Full-text available
  • Jan 2022
Introduction. Monechma ciliatum L. seeds are rich in proteins, carbohydrates, oils and mineral contents. Researchers have focused on new production development but there is no available data on the impact of processing techniques on the quality of the seeds. Our study aimed to investigate the impact of boiling, roasting, and germination on the composition and nutritional value of Monechma ciliatum (black mahlab) seeds. Study objects and methods. We analyzed 7 kg of black mahlab seeds purchased from the local market. We applied standard methods used in boiling, roasting, and germination techniques. Proximate analyses were performed using the methods of the Association of Official Analytical Chemists. Minerals were analyzed by inductively coupled plasma-mass spectrometry (ICP-MS), and fatty acids were determined by gas chromatography. Tocopherols and amino acids in processed seeds were determined by high-performance liquid chromatography. Results and discussion. The results showed that the proximate compositions of untreated, boiled, roasted, and germinated mahlab seeds were affected by boiling, roasting, and germination techniques. Most of the nutritional values were enhanced by all the treatments. In particular, all the processing techniques increased the protein content. Boiling and roasting increased the fat content, while boiling and germination increased the fiber content. Tocopherols were higher only in the germinated samples. Amino acids were increased by all the techniques. Minerals were affected by all the techniques, except for Na, which was higher in the germinated sample. Conclusion. Boiling, roasting, and germination enhanced significantly the chemical composition of Monechma ciliatum seeds, which make them a value ingredient to develop new food products.
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Goyder 2023 New species from Angola PhytoKeys
Article
Full-text available
  • Sep 2023
Four new species are described from central and eastern Angola and adjacent NW Zambia. All occur in Kalahari sand savannas rich in endemic and more widely distributed geoxylic suffrutices. Despite being known from very few collections, the conservation status of one of these new species is assessed as Least Concern, as these grasslands are nutrient-poor, are in remote sparsely populated areas, and are not threatened with conversion to agriculture. The remaining three are treated as Data Deficient. In addition, one new combination is provided for Ancylanthos rubiginosus Desf. under Vangueria as V. rubiginosa (Desf.) Lantz is an illegitimate later homonym. We also make orthographic corrections to specific epithets commemorating Ilse von Nolde, a collector who made important collections from Quela in Malange in the 1930s.
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A taxonomic revision of Acanthaceae tribe Barlerieae in Angola and Namibia. Part 1
Article
Full-text available
  • Mar 2019
In the first part of a taxonomic revision of the tribe Barlerieae (Acanthaceae) in Angola and Namibia, two of the four major clades of the genus Barleria L. are studied in detail, namely subg. Barleria and subg. Prionitis sect. Prionitis. Using standard morphology-based taxonomic methods, a total of 32 species and 39 taxa are delimited, of which 12 are endemic to Namibia, nine are endemic to Angola, a further four are restricted to the two countries combined and four are largely confined to the region. Hence, nearly 70% of the taxa are endemic or near-endemic to Angola and Namibia. Full descriptions, citations of types and representative specimens, and notes on their habitat(s), ecology and biogeography are provided for each taxon. Seven new species (Barleria calole, B. crabbeoides, B. craveniae, B. deserticola, B. fernandae, B. namba and B. rupicola), two new subspecies (B. lanceolata subsp. tristis and B. prionitoides subsp. strigulosa) and one new variety (B. rigida var. spinissima) are described. In addition, Barleria namutonensis Oberm. is treated as a subspecies of B. prionitoides, and B. dinteri Oberm. is treated as a subspecies of B. coriacea Oberm. and the necessary new combinations are made, whilst B. schenckii Schinz is resurrected as a good species. Eleven names in Barleria are lectotypified. A preliminary assessment of the conservation status and extinction risk is provided for each taxon. Of the 39 taxa enumerated, the majority are assessed to be of Least Concern, but one is assessed as Endangered, four as Vulnerable, one as Near Threatened and six are currently considered to be Data Deficient.
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A synopsis of Rhinacanthus (Acanthaceae) in Angola and Namibia
Article
Full-text available
  • Jun 2018
The three species of the genus Rhinacanthus Nees occurring in Angola and Namibia are documented, including a full description of the new species R. angolensis I. Darbysh. and an expanded description of the scarce species R. kaokoensis K. Balkwill & S. D. Will. A key to their identification is provided, together with notes on their conservation status and species affinities.
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RADseq dataset with 90% missing data fully resolves recent radiation of Petalidium (Acanthaceae) in the ultra‐arid deserts of Namibia
Article
Full-text available
  • Aug 2017
Deserts, even those at tropical latitudes, often have strikingly low levels of plant diversity, particularly within genera. One remarkable exception to this pattern is the genus Petalidium (Acanthaceae), in which 37 of 40 named species occupy one of the driest environments on Earth, the Namib Desert of Namibia and neighboring Angola. To contribute to understanding this enigmatic diversity, we generated RADseq data for 47 accessions of Petalidium representing 22 species. We explored the impacts of 18 different combinations of assembly parameters in de novo assembly of the data across nine levels of missing data plus a best practice assembly using a reference Acanthaceae genome for a total of 171 sequence datasets assembled. RADseq data assembled at several thresholds of missing data, including 90% missing data, yielded phylogenetic hypotheses of Petalidium that were confidently and nearly fully resolved, which is notable given that divergence time analyses suggest a crown age for African species of 3.6–1.4 Ma. De novo assembly of our data yielded the most strongly supported and well-resolved topologies; in contrast, reference-based assembly performed poorly, perhaps due in part to moderate phylogenetic divergence between the reference genome, Ruellia speciosa, and the ingroup. Overall, we found that Petalidium, despite the harshness of the environment in which species occur, shows a net diversification rate (0.8–2.1 species per my) on par with those of diverse genera in tropical, Mediterranean, and alpine environments.
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Unraveling relationships in the morphologically diverse and taxonomically challenging “justicioid” lineage (Acanthaceae: Justicieae)
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Full-text available
  • Jun 2017
This is the first study in Acanthaceae to provide a robust and well-sampled phylogenetic framework for the ‘justicioid’ lineage (Acanthaceae: Acanthoideae: Justicieae), a clade that includes at least 1000 species that together occupy essentially the entire distributional range of the family as a whole. Species are morphologically diverse and occupy a wide variety of habitats. Although much progress has been made in understanding phylogenetic relationships within Acanthaceae, well-sampled phylogenetic studies are lacking for ‘justicioids’. DNA sequence data from six regions for 178 accessions from across the entire lineage were used to test monophyly and validity of 17 putative genera of ‘justicioids’ on a global scale. Included among these genera is the species-rich genus Justicia for which we evaluate the current morphology-based classification with a focus on Old World taxa. Our results strongly support monophyly of New World ‘justicioids’, including all New World species of Justicia plus species assigned to at least five other smaller genera. Further, with a greatly expanded taxon sample, we confirm our earlier unexpected result that Diclipterinae are embedded within ‘justicioids.’ As traditionally circumscribed, Diclipterinae lack the morphological traits that mark ‘justicioids’ and thus had not previously been associated with that group. This study further extends this unexpected result by placing a number of Old World ‘justicioids’ in a clade that is sister to Diclipterinae. Although some genera of ‘justicioids,’ notably of Diclipterinae, are monophyletic, Justicia is grossly polyphyletic. We explore these results by examining the diagnostic utility of morphological characters for major clades with a focus on inflorescences, androecia, pollen and seeds. We further use our results as context for documenting evolutionary patterns of floral traits and pollinator relationships, as well as biogeographic patterns, across this species-rich and diverse clade.
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A brief botanical survey into Kumbira forest, an isolated patch of Guineo-Congolian biome
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  • Jun 2016
Kumbira forest is a discrete patch of moist forest of Guineo-Congolian biome in Western Angola central scarp and runs through Cuanza Norte and Cuanza Sul province. The project aimed to document the floristic diversity of the Angolan escarpment, a combination of general walk-over survey, plant specimen collection and sight observation was used to aid the characterization of the vegetation. Over 100 plant specimens in flower or fruit were collected within four identified vegetation types. The list of species includes two new records of Guineo-Congolian species in Angola, one new record for the country and one potential new species.
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Documenting Biogeographical Patterns of African Timber Species Using Herbarium Records: A Conservation Perspective Based on Native Trees from Angola
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In many tropical regions the development of informed conservation strategies is hindered by a dearth of biodiversity information. Biological collections can help to overcome this problem, by providing baseline information to guide research and conservation efforts. This study focuses on the timber trees of Angola, combining herbarium (2670 records) and bibliographic data to identify the main timber species, document biogeographic patterns and identify conservation priorities. The study recognized 18 key species, most of which are threatened or near-threatened globally, or lack formal conservation assessments. Biogeographical analysis reveals three groups of species associated with the enclave of Cabinda and northwest Angola, which occur primarily in Guineo-Congolian rainforests, and evergreen forests and woodlands. The fourth group is widespread across the country, and is mostly associated with dry forests. There is little correspondence between the spatial pattern of species groups and the ecoregions adopted by WWF, suggesting that these may not provide an adequate basis for conservation planning for Angolan timber trees. Eight of the species evaluated should be given high conservation priority since they are of global conservation concern, they have very restricted distributions in Angola, their historical collection localities are largely outside protected areas and they may be under increasing logging pressure. High conservation priority was also attributed to another three species that have a large proportion of their global range concentrated in Angola and that occur in dry forests where deforestation rates are high. Our results suggest that timber tree species in Angola may be under increasing risk, thus calling for efforts to promote their conservation and sustainable exploitation. The study also highlights the importance of studying historic herbarium collections in poorly explored regions of the tropics, though new field surveys remain a priority to update historical information.
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The genus Hypoestes (Acanthaceae) in Angola
Article
  • Dec 2015
A synoptic account of the genus Hypoestes (Acanthaceae) in Angola is presented, including a key to the four species known from the country. The new species Hypoestes ecbolioides I. Darbysh. is described and illustrated from southwest Angola and its affinities and conservation status are discussed.
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Two new species in Justicia sect. Harnieria (Acanthaceae) from tropical Africa
Article
  • Jun 2008
  • NORD J BOT
Two new species of Justicia sect. Harnieria (Acanthaceae) are described: J. afromontana from Sudan and Uganda, and J. huambensis from Angola.
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