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Baldwin & Wood • Systematics and biogeography of the Bahia alliance
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INTRODUCTION
Lepidium
Microseris
Apostates
Pacifigeron
and Metatrophis
Miocalles
Origin of the Rapa endemic genus Apostates: Revisiting major
disjunctions and evolutionary conservatism in the Bahia alliance
(Compositae: Bahieae)
Bruce G. Baldwin & Kenneth R. Wood
1 Jepson Herbarium and Department of Integrative Biology, University of California, Berkeley, California 94720, U.S.A.
2 National Tropical Botanical Garden, Kalaheo, Hawaii 96741, U.S.A.
Bruce G. Baldwin, bbaldwin@berkeley.edu
ORCID
DOI 655
Abstract
Apostates
Bahia
Apostates
Bahia
Bahia
Bahia
Apostates
Keywords
Supplementary Material
Baldwin & Wood • Systematics and biogeography of the Bahia alliance
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VeronicaHebe
Corokia
Astelia
Apostates
A.M. Stokes 186
Olearia O. rapae
Olearia
O. rapaeOlearia
O. rapaeOlearia
O. rapae
Apostates
Apostates
Apostates
Apostates
Apostates on a
ndhFtrnK
rpl16 psbAtrnH Apostates
Apostates
Apostates
Apostates
Fig. .Apostates rapae(lef t),
(right).
Baldwin & Wood • Systematics and biogeography of the Bahia alliance
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Fig. .BahiaA, Bahia neomexicana (= Picradeniopsis multif lora
B, Bahia absinthifolia (= Picradeniopsis absinthifoliaC, Bahia
ambrosioidesD,Apostates rapaeE, Apostates rapae
F, Hymenothrix wrightiiG, Amauriopsis dissecta (= Hymenothrix dissecta
H, Palafoxia arida
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MATERIALS AND METHODS
Taxon sampling. —
Apostates rapae col
Wood &
Bahia Bahia
Bahia
Apostates
The families and genera of vascular plants
AchyropappusAmauriopsis ApostatesBahia
Cephalobembix Picradeniopsis
Florestina Holoschkuhria
Hymenothrix Nothoschkuhria Palafoxia
PlatyschkuhriaSchkuhria
Chaetymenia
Bahia
Picradeniopsis
B. pringlei
B. neomexicana
in Schkuhria S. multiflora
Amauriopsis Bahia
A. janakosana
Achyropappus
A. depauperatus
ApostatesHoloschkuhriaNothoschkuhria
and Platy schkuhriaHymenothrix
H. greenmanii
Schkuhria
S. greenmaniiHymenothrix greenmanii
Florestina
Palafoxia
DNA extraction and sequencing. —
ndhFndhF-ycf1 inter
trnK intron and 5
matK
rpl16
psbAtrnH
Analyses of phylogeny, biogeography, and morphological
character evolution. —
Bahia pedata
matKndhFrbcL
Amauriopsis dissectaBahia
absinthifoliaChaetymenia peduncularis
Florestina pedata Palafoxia arida
Peucephyllum schottii
Dimeresia howellii Chaenactis
douglasii C. santolinoides
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Bahia
Bahia al
Schkuhria
Barnedesia
Dasyphyllum
Bahia
Bahia
Bahia
RESULTS
Phylogenetic resolution, congruence, and incongruence.
—Bahia
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Amauriopsis Hymenothrix Platy-
schkuhria
Bahia
matKndhFrbcL
Bahia
Schkuhria
S. degenericaNothoschkuhria
C. pedunculata B978
S. wislizeni B974
S. schkuhrioides nx3/cPOP-150
S. wrightii K19020
S. pinnata nx2/cx3
Ac. anthemoides K15420
Ac. queretarensis S8344
B. ambrosioides C4574
Ho. tetramera nx2/cC3654
N. degenerica B7991
Apostates rapae x2
B. woodhousei nx2/cH8652
B. absinthifolia x2
B. oppositifolia H3153
B. neomexicana nx3/cH23490
B. bigelovii nx3/cP6384ind2
B. xylopoda nx3/cN28852
N32946
(B. schaffneri nHB6388/c-)
F. platyphylla K15532
F. tripteris B970
Pa. arida B784
Pa. texana B969
Pl. integrifolia nx2/cx1
Am. autumnalis x2
Am. biternata x2
Am. glandulosa A326
Hy. loomisii / wrightii
Hy. wislizeni x2
Am. dissecta x2
Am. dissecta I122
Hy. palmeri x2
(Am. pedata n-/cS438)
Chaetymenia (outgroup) Chaetymenia (outgroup)
Schkuhria
Schkuhria
Achyropappus
Achyropappus
Bahia Bahia
Holoschkuhria Holoschkuhria
Nothoschkuhria Nothoschkuhria
Apostates Apostates
Picradeniopsis
Picradeniopsis
Florestina
Florestina
Palafoxia
Palafoxia
Platyschkuhria
Platyschkuhria
Hymenothrix
Hymenothrix
1.00/100
1.00/84
1.00/100
1.00/100
1.00/67
1.00/95
1.00/98
1.00/99
1.00/100
1.00/100 1.00/100
1.00/100
1.00/100
0.97/80
1.00/100
1.00/100
1.00/100
1.00/99
1.00/100
1.00/100
1.00
1.00
1.00/87
1.00/100
0.99/95
1.00/100
1.00/100
1.00/95
1.00/99
1.00/100
1.00/100
1.00/92
0.99/90
1.00
1/100
0.98/93
1.00/98
1.00/96 1.00/100
1.00/100
1.00/100
1.00/98 1.00/100
1.00/100
1.00/99
97
100
97
1
1
2
2
3
3
4
4
a
b
c
a
b
c
1.00/100
Fig. . (lef t) (right) Bahia
Ac AchyropappusAm Amauriopsis B. Bahia C.ChaetymeniaF. Florestina Ho. HoloschkuhriaHy.
Hymenothrix NNothoschkuhria Pa. Palafoxia Pl. PlatyschkuhriaS.Schkuhria.
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degenericaS. multiflora (= Bahia
neomexicanaAchyropappus Bahia am-
brosioides HoloschkuhriaApostates
Bahia
B. neomexicanaNothoschkuhriaAmauriopsis
FlorestinaHymeno-
thrix Palafoxia
Platyschkuhria
Flores-
tinaPalafoxia (c) AmauriopsisHymenothrix
PlatyschkuhriaBahia
B. neomexicana
Bahia
absinthifolia B. oppositifolia
B. bigelovii B. neomexi-
canaB. schaffneri
B. xylopoda Schkuhria
S. pinnata
S. wrightii
can S. wislizeni S. schkuhrioides
BahiaB. neomexi-
canaApostatesAmauriopsis dissecta
Hymenothrix palmeri
Achyropappus Bahia ambrosi-
oides Holoschkuhria
B. ambrosioidesHoloschkuhria
AchyropappusB. ambrosioides
Fig. .Bahia
Bahia
4
AcAchyropappusAmAmauriopsis B. BahiaC.Chaetymenia
F. Florestina Ho. HoloschkuhriaHy. Hymenothrix N.Nothoschkuhria Pa. Palafoxia Pl. Platyschkuhria S.Schkuhria.
051 01 52 02 53 0
Apostates rapae (2)
B. xylopoda (2)
Pl. integrifolia B938(cl1)
Ac. anthemoides K15420
B. neomexicana F5658
S. wrightii K19020
B. absinthifolia S10714
B. schaffneri HB6388
Hy. loomisii/wrightii (1/1)
B. woodhousei (2)
Pa. arida B784
F. tripteris B970
C. peduncularis B978
Ho. tetramera C3654
Am. biternata VD93-789
Ho. tetramera O90733
B. absinthifolia B967
Pl. integrifolia B938(cl2)
Hy. palmeri RG97-1158
S. pinnata C103-03
B. bigelovii P6384(2 ind)
Am. biternata D1037
S. schkuhrioides (3)
Hy. palmeri S2722
N. degenerica B7991
Ac. queretarensis S8344
Am. autumnalis V7972
Pa. texana B969
Am. autumnalis V6604
B. oppositifolia H3153
B. neomexicana (2)
B. ambrosioides C4574
Am. dissecta (2)
F. platyphylla K15532
S. pinnata C111-07
Hy. wislizeni SJB s.n.
B. bigelovii P6384(1 ind)
S. wislizeni B974
Am. dissecta I122
Am. glandulosa A326
B. xylopoda GP1452
Hy. wislizeni C1517
Alternate leaves
Alternate leaves
(Opposite leaves)
1/100
1/100
1/100
0.98/93
1/100
1/100 1/99
1/100
*1/97
1/100
1/100
1/87
0.99/95
1/100
1/100
1/95
1/100
1/99
*1/92
1/100
-/71
0.99/90
*1/98
1/96
0.89/-
1/100
1/98
1/100
1/100
-/53
1/99
-/-
0.93/66
1/100
1/100
1/100
-/-
-/-
South America
North America
Rapa
Biogeography
(clade color):
:
Head type
(taxon label color):
Radiate heads
Discoid heads
Ma
1/96
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Florestina Palafoxia Amauriopsis
HymenopappusPlatyschkuhria
Florestina Palafoxia
Palafoxia AmauriopsisHymenothrixPlatyschkuhria
Bahia bigeloviiB. neomexicana
B. xylopodaB. bigelovii
B. xylopodaB. schaffneri
B. neomexicanaB. bigelovii
B. neomexicanaB. xylopoda
Biogeographic model tests and estimates. —
Bahia
Morphological evolution. —
AmauriopsisHymenothrixPlatyschkuhria and once on
Nothoschkuhria
Palafoxia
Bahia alliance
Palafoxia
DISCUSSION
Apostates belongs t o the Bahia alliance. —
Apos-
tates
Apostates
Apostates
Apostates
Bahia
Schkuhria
Bahia
Bahia Apostates
Bahia
B. neomexicana Notho schkuhria
Bahia
BahiaB. ambrosioides
Achyropappus
Holoschkuhria
Bahia
BahiaAmauriopsis
FlorestinaHymenothrix
PalafoxiaPlatyschkuhria
Apostates represents long-distance dispersal from North
America. — Apostates
Hypericophyllum Apos-
tates
Apostates
Apostates
Apostates
ApostatesBahia alliance cannot
Tubulifloridites lilliei
Other major disjunctions in the Bahia alliance. —
Apostates
Tab le .
L
Baldwin & Wood • Systematics and biogeography of the Bahia alliance
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Bahia Apostates
Bahia
Apostates and
HoloschkuhriaBahia
Apostates
Bahia
Apostates
Holoschkuhria
Nothoschkuhria
Bahia neomexicana
Bahia ambrosioidesHoloschkuhria
Bahia ambrosioides and Holoschkuhria
Bahia ambrosioides
Holoschkuhria (H. tetramera
Bahia
B. ambrosioidesAchyropap-
pus
Achyropappus
B. ambrosioidesHoloschkuhria
in B. neomexicana (= Schkuhria multiflora
Schkuhria
Bahia al
Nothoschkuhria
Bahia
B. neomexicana and Apostates
Comparable biogeographic patterns in the southern Poly-
nesian flora. —
Apostates
Fitchia
Oparanthus
Narvalina Selleophytum
Petrobium
FitchiaOparanthus
Bahia Plantago
P. rapensis P. rupicola
P. macrocarpa
Metrosideros
Pittosporum
Plantago
Nesoluma
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Melicope
2014) and Santalum
Intrinsic and extrinsic dispersal factors. —
Apos-
tates
Bahia
Bahia
Amauriopsis (= Bahia
Apostates
Apostates
Phyllanthus
Epicephala
Evolutionary conservatism in the Bahia alliance? —
Bahia
acter in Bahia
Amauriopsis
Ba-
hia
Amauriopsis (= Bahia
Hymenothrix and
Platy schkuhria Bahia
B. neomexicana (= Schkuhria mul-
tifloraApostates
Bahia
Nothoschkuhria Amauri-
opsisHymenothrixPlatyschkuhria
Bahia
Bahia
Apostates
Hymenothrix Palafoxia
AchyropappusAmauriopsisPlatyschkuhria
Schkuhria.
Amauriopsis
Bahia
Achyropappus and Platyschkuhria
BahiaB. neo-
mexicana Bahia
Schkuhria S. multiflora
Schkuhria
Taxonomic treatment. —
Bahia
Florestina and
Palafoxia
Platyschkuhria
AmauriopsisHymenothrixNotho-
schkuhriaSchkuhria degenerica
Schkuhria
Amauriopsis (= Bahia
Hymenothrix
Amauriopsis and Hymenothrix
Bahia alliance
Hymenothrix
Bahia Apostates
BahiaPic-
radeniopsisNothoschkuhriaAchyropappusB. am-
brosioidesHoloschkuhria
Hymenothrix
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Bahia (B. ambrosioides
Apostates
Picradeniopsis
Cephalobembixx B. bigeloviiB. neo-
mexicanaB. pringlei
B. xylopoda B. schaffneriB. xylopoda
Picradeniopsis x B. absinthifoliaB. oppositi-
foliaB. woodhousei
to Apostates
B. woodhousei
B. absinthifolia B. oppositifolia
B. woodhouseiPicradeniop-
sisP. oppositi foliaBahia oppositi-
folia) and P. woodhouseiBahia woodhousei)
x
Achyropappus
ApostatesBahia
Holoschkuhria
Key to genera of the Bahia alliance
2
4
Nothoschkuhria
Platyschkuhria
Hymenothrix
5
6
Florestina
Palafoxia
Schkuhria
8
Apostates
Holoschkuhria
Achyropappus
10
Bahia
Picradeniopsis
New combinations for the proposed taxonomic changes
Hymenothrix autumnaliscomb.
nov.Bahia autumnalis
Amauriopsis autumnalis
M.C . Johnston & J. Crutchfield 5856
Hymenothrix biternata c omb . nov.
Bahia biternata
Schkuhria biternata
Eriophyllum biternatum
Villanova biternata
Amaur -
iopsis biternata
C. Wright 1256 (GH
Hymenothrix dissecta comb. nov.
Amauria? dissecta
Villanova chrysanthemoides
V. dissectaBahia
chrysanthemoides
Bahia dissecta
Eriophyllum chrysanthemoides
Villanova dis-
secta
Amauriopsis dissecta
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A. Fendler 537
= Bahia dissecta anisopappa
I.L. Wiggins &
D. Demaree 4870
Hymenothrix glandulopubescens
comb. nov.Hymenopappus glandulopubescens
Waterfall
12576
= Bahia glandulosa
Hymenothrix glandulosa
Amauriopsis glan-
dulosa
2010 – Lectotype, designated here
E. Palmer 492
(E. Palmer 492
C.G. Pringle 308Bahia
glandulosa
Palmer 492
Hymenothrix janakosanacomb.
nov.Amauriopsis janakosana
F. Chiang, T. Wendt & M.C. John-
ston 9665
Hymenothrix pedata comb. nov.
Bahia pedata
Schkuhria pedata
Eriophyl-
lum pedatum
Amauriopsis pedata
C. Wright 387 (GH
Picradeniopsis absinthifolia comb.
nov. Bahia absinthifolia
Eriophyllum absinthifolium
Ha rt weg 128
= Bahia dealbata
Bahia absinthifolia deal
-
bata
Picradeniopsis dealbata
Lecto type (designated here):
Dr. Gregg s.n. (GH
Hart weg 128
Bahia absinthifolia
Hartweg 128
B. de-
albata
Picradeniopsis bigelovii com b. nov.
Bahia bigelovii
Schkuhria bigelovii
Eriophyllum
bigelovii
Bigelow s.n.
Picradeniopsis multiflora
comb. nov.Schkuhria multiflora
Achyropappus
schkuhrioides
Achyropappus schkuhrioides
Bahia gil-
liesii
Bahia multiflora
Gillies 147
= Schkuhria neomexicana
Neo-MexicanaAm-
blyopappus neomexicanus
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Bahia neomexicana
Achyropappus neomexicanus
Cephalobembix
neomexicana
A. Fendler 458 (GH
= Schkuhria pusilla
Rothia pusilla
Schkuhria multif lorapusilla
d’Orbigny 1403
= Rothia intermedia
Lectotype (designated here):
O. Kuntze s.n.
= Schkuhria pusilla aristata
Schkuhria multi-
flora aristata
R.E. Fries 812b
= Schkuhria pusilla longipedicellata
longepedicellata
Bodenbender s.n.
Rothia intermedia
Picradeniopsis pringleicomb. nov.
Bahia pringlei
C.G. Pringle
6407
Picradeniopsis schaffneri comb.
nov.Bahia schaffneri
C.G. Pringle
3028
Picradeniopsis schaffneriaristata
co mb. nov. Bahia aristata
Bahia schaffneriaristata
C.G. Pringle 5127 (GH barcode
Picradeniopsis xylopoda com b. n ov.
Bahia xylopoda
C.G. Pringle 6931
= Bahia ehrenbergii
Ehrenberg
362
Extreme endangerment of Apostates. —
Apostates
rapae
A. rapae
Myoporum rapense Scaevola taccada
tuamotuensis Eugenia reinwardtiana
and Osteomeles anthyllidifolia
Dianella intermedia punctata Portulaca
lutea and Lycium sandwicense
Cyperus javanicus Chrysopogon
aciculatus Ischaemum stokesii
Asplenium obtusatum and Polystichum
rapense
A. rapae
Psidium cattleianum Miscanthus florid-
ulus, Scirpus nodosus Sporobolus
elongatus and Eleusine indica
A. rapae
Baldwin & Wood • Systematics and biogeography of the Bahia alliance
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A. rapae
A. rapae
Apostates
Apos-
tates
A. rapae
ACKNOWLEDGMENTS
Apostates
rapae
Appelhans, M.S., Wen, J, Wood, K.R., Allan G.J., Zimmer, E.A. &
Wagner, W.L.
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