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Generic circumscription and relationships of southern ...
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Generic circumscription and relationships of southern African representatives of Hypoxis and allies (Hypoxidaceae, Asparagales)
Sanele Shiba, Michelle van der BankTheAfricanCentreforDNABarcoding(ACDB),DepartmentofBotany&PlantBiotechnology,UniversityofJohannesburg,SouthAfrica
ABSTRACTBackground. The popularity of Hypoxis L. as a medicinal plant has resulted in unsustainable harvesting practices of rhizomes from the wild. This exploitation has accelerated theneed for correct species names and circumscriptions. However, species delimitation in Hypoxis is problematic and despite several attempts, the systematics of the genus remainslargely unresolved. This is mainly due to the lack of distinct morphological boundaries separating species. Here, we explore the generic circumscriptions of Hypoxis and alliedgenera within Hypoxidaceae using five plastid DNA regions (rbcLa, matK, trnL-F, ycf1 and trnS-G). Results. Findings from our study indicate that Hypoxis is not monophyletic andis represented by at least three distinct lineages. Using the phylogeny produced from the study Hypoxis samples sold at traditional medicinal markets in South Africa could besuccessfully identify. Significance. We proposed to transfer Rhodohypoxis Nel back into Hypoxis as well as the newly described genus Sinocurculigo from China to CurculigoGaertn.
Hypoxis L. represents the largest genus in theHypoxidaceae R. Br. with about 90 species (Kocyan et al.,2011).
The popularity of Hypoxis as a medicinal plant has resultedin unsustainable harvesting practices of rhizomes from thewild. This exploitation has accelerated the need for correctspecies names and circumscriptions (Zimudzi & Kativu,2006; Singh, 2009).
However, species delimitation in Hypoxis is problematicand despite several attempts, the systematics of the genusremains largely unresolved. This is mainly due to the lackof distinct morphological boundaries separating species(Wiland-Szymanska, 2009).
Aim: To resolve the phylogenetic relationships withinHypoxidaceae as well as producing a species levelphylogeny for the approximately 90 species of the genusHypoxis.
Doyle, J.J. & Doyle, J.L., 1987. A rapid isolation procedure for small amounts of leaf tissue. PhytochemicalBulletin 19: 11-15.
Swofford, D. L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. SinauerAssociates, Sunderland, MA.
Kocyan, A., Snijman, D.A., Forest, F., Devey, D.S., Freudenstein, J. V., Wiland-Szymańska, J., Chase, M.W. &Rudall, P.J. 2011. Molecular phylogenetics of Hypoxidaceae - Evidence from plastid DNA data andinferences on morphology and biogeography. Molecular Phylogenetics and Evolution. 60(1):122-136. DOI:10.1016/j.ympev.2011.02.021.
Kocyan, A. & Wiland-Szymanska, J. 2016. Friedmannia: a new genus from the Seychelles and the beginning of ageneric realignment of Curculigo (Hypoxidaceae). Phytotaxa. 283(1): 54-64.
Snijman, D.A. & Kocyan, A. (2013) The genus Pauridia (Hypoxidaceae) amplified to include Hypoxis sect. Ianthe,Saniella and Spiloxene, with revised nomenclature and typification. Phytotaxa 116: 19-33.
Wiland-Szymanska, J. 2009. The Genus Hypoxis L. (Hypoxidaceae) in the East Tropical Africa: variability,distribution and conservation status. Biodiv. Res. Conserv. 14: 1-129.
Zimudzi, C. & Kativu, S. 2006. The African potato (Hypoxis hemerocallidea): identity and status in Zimbabwe.Kirkia. 18(pt. 2): 175-176.
MATERIAL & METHODSSamplingA total of 93 taxa were included in the data set. Sampleswere either collected in the field, sourced from herbariumvouchers or downloaded from GenBank.
DNA extraction, amplification and sequencingDNA was extracted using the 10X CTAB method of Doyle& Doyle (1987). Standard protocols were followed for PCRand sequencing, using three plastid regions (rbcLa, trnL-Fand trnS-G).
Data analysisA phylogenetic tree was generated using MaximumParsimony (MP) in PAUP* (Swofford, 2002).
INTRODUCTION
FIGURE 1: Representatives of the southern African genera of Hypoxidaceae
RESULTS & DISCUSSION
FIGURE 2: Phylogenetic tree for the family Hypoxidaceae based on the combined data set showing four well supported clades.
REFERENCES
ACKNOWLEDGEMENTS
CONCLUSIONAlthough the backbone of the phylogeny was not resolved, four well supportedclades could be identified (Clades A – D). Based on the results we propose totransfer Rhodohypoxis back into Hypoxis as well as the newly described genusSinocurculigo to Curculigo.
In Figure 2, there is a 100% support for the monophyly of the familyHypoxidaceae, with four major clades being identified.
There is 100% support for Clade A. The genus Rhodohypoxis was established in1914 to accommodate the Hypoxis plants with white, pink or red flowers. The onlyother morphological difference between Rodohypoxis and Hypoxis is theconcealed stamens and style in Rhodohypoxis.
Within Clade B (99%), Molineria is nested within Curculigo. Since Curculigo hasnomenclatural priority, Molineria species should be transferred to Curculigo withCurculigo seychellensis being the exception. Clade B also suggests that thenewly described genus Sinocurculigo from China should be transferred to thegenus Curculigo. Curculigo seychellensis forms a monophyletic Seychelles cladewith Hypoxidia maheensis and Hypoxidia rhizophylla. A new monotypic genus(Friedmannia), endemic to the Seychelles was proposed to accommodate thesespecies (Kocyan & Wiland-Szymanska, 2016).
There is a high support (100%) for the monophyly of the Empodium species(Clade C) included in this study. There is a high support (85%) for the cladecontaining Pauridia, Spiloxene, Saniella and Hypoxis section Ianthe (Clade D).This supports findings by Snijman and Kocyan (2013) who transferred all thespecies of Spiloxene, Saniella and Hypoxis section Ianthe to Pauridia.
Hypoxishirsuta Pauridia aquatica
Saniella verna Spiloxene capensis
Empodium gloriosum Rhodohypoxisbaurii
Astelia alpina KocyanAK981010 1 01 Chase1103Astelia banksii Chase1072Blandfordia nobilis Chase2835Blandfordia punicea AK981013 1 01 Chase519
Curculigo erecta Leiden932789
Curculigo racemosa Leiden970647
Molineria latifolia KocyanAK981023 1 03
Curculigo finlaysoniana CookRixSchneller23
Curculigo orchioides Janarthanam TCMK696
Curculigo pilosa subsp major Ehrich266
Curculigo sinensis KunmingBotanicalGarden
Molineria crassifolia KunmingBotanicalGarden
Molineria capitulata KocyanAK981228 1 01
Sinocurculigo taishanica
Molineria aff latifolia BGM
unich98 3407 Bogner2650
Molineria villosa KocyanAK971031 1 03
Curculigo seychellensis Bonn
Empodium
elongatum Snijm
an1908
Em
podium sp. D
esmet2979
Em
podium flexile S
nijman1706E
mpo
dium
plic
atum
BG
Mun
ich
Em
podi
um v
erat
rifol
ium
Koc
yanA
K99
0309
1 0
3
Hyp
oxis
acu
min
ata
BNR
H00
0316
1
Hyp
oxis
acu
min
ata
BT13
09
Hyp
oxis
acu
min
ata
BT13
07
Hyp
oxis
ang
ustif
olia
var
. ang
astif
olia
BN
RH
0006
307
Hypo
xis c
osta
ta B
NRH0
0092
68
Hypo
xis h
emer
ocal
lidea
BNR
H000
3965
Hypo
xis in
terje
cta B
NRH0
0091
43
Hypox
is kra
ussia
na B
NRH0006
312
Hypox
is lon
gifoli
a BNRH00
0872
5
Hypox
is lon
gifoli
a BNRH00
0629
7
Hypoxis
obtusa BNRH0003269
Hypoxis parvif
olia BNRH0009285
Hypoxis rigidula var. ri
gidula BNRH0006261
Hypoxis rigidula var. ri
gidula BNRH0006262
Hypoxis colchicifolia BNRH0006375
Hypoxis costata BT1306
Hypoxis galpinii BNRH0006328Hypoxis galpinii BT1305
Hypoxis rigidula var. pilosissima BNRH0006534Hypoxis hemerocallidea BGBerlin635807Hypoxis hemerocallidea BNRH0006241
Hypoxis rigidula cf. rigidula BB14828
Hypoxis parvifolia BNRH0006219
Hypoxis sp. Bonn126
Hypoxis angustifolia KewGardens1973 2957
Hypoxis angustifolia var. buchananii BNRH0009208
Hypoxis argentea BNRH0006257
Hypoxis argentea var. argentea BNRH0002746
Hypoxis aurea KocyanAK426
Hypoxis curtissii Chase108
Hypoxis hirsuta Wipf2209
Hypoxis juncea WhittenK
Hypoxis decumbens KewGardens1976 6038
Hypoxis filiformis Basel KocyanAK438
Hypoxis filiformis BNRH0009185
Hypoxis hygrometrica BGBerlin43336
Hypoxis limicola BNRH0006314
Hypoxis limicola BT1311
Rodohypoxis baurii var. baurii BSP10497
Rodohypoxis baurii var. confecta BN
RH
0023126H
ypoxis parvula NH
556
Rhodohypoxis baurii BG
Basel KocyanAK990522 1 02R
hodohypoxis baurii var. baurii Singh554
Hypoxis setosa ZS
S29 156
Hyp
oxis
vill
osa
Kew
Gar
dens
1972
394
7H
ypox
is g
labe
lla Y
1498
9H
ypox
is o
ccid
enta
lis K
ew22
34B
Sani
ella
occ
iden
tali
Snijm
an20
59Sa
niel
la v
erna
BG
Edin
burg
h199
7 32
02
Spilo
xene
cap
ensi
s Ar
b728
Spilo
xene
alb
a Sn
ijman
2106
Spilo
xene
aqu
atica
Sni
jman
2113
Spilo
xene
mon
ophy
lla P
ater
son
Jone
s913
Spilo
xene
nan
a Sn
ijman
1865
a
Spilox
ene p
usilla
Snij
man18
60
Spilox
ene s
chlec
hteri P
arker4
69
Paurid
ia lon
gitub
a Koc
yanA
K9806
24 1
01
Pauridia longituba Snijm
an1440
Pauridia minuta Snijman1812
Spiloxene flaccida Goldblatt Manning9585
Spiloxene trifurcillata McMaster208084
Spiloxene linearis Snijman1754
Spiloxene minuta Parker452
Spiloxene scullyi Harrower3271
Lanaria lanata Orchard342
Milligania stylosa Kocyan981011 3 01 Chase511
Collospermum hastatum BG875 767
5289100
95
96
100
99
99
85 92
62
61 65
82
95
100100
100
916210
0
100
86
58
79
72
59
94
72
50 87
85
86
55
6886
53
9377
9674
Bootstrap consensus tree
Curculigo scorzonerifolia Gorts et al. 449
Curculigo seychellensis Kuffer 26 11 2002
Hypoxidia m
aheensis Kuffer 26 11 2002
Hypoxidia rhizophylla Kuffer 26 11 2002
Clade A
Clade CClade B
Clade D
HypoxidaceaeKEY
PosterID741