Table Of ContentCLASSIFICATION, ORIGIN,
Michael
Steven WagsldJ]/ Bayly,'
/.
J.
AND Philip Carnoch-Jones,^
OF
DIVERSIFICATION
./.
and
Dirk Alharlr^
C,
NEW
THE ZEALAND HEBES
(SCROPHULARIACEAE)'
AlJSTKACT
NfU
Zealand lu-hcs (Srn»f)luilariat'eae) arr nu'inlu'is of a lar^c Soiillu'iii Irrnisplu're clade nested williin Ve-
Till'
I
ronica. Analysis of IIS and rlnX. scijuences siit!;gfsls Hiat the Nt'w Zealand species are derived from a single eonunon
Nen
fonnder population
ancesh)! that arrived via long-distance dispersal. After the estahlishnient of iliis initial in
Zealand, hehes have undergone least two major episod«»s of diversification, giving rise to six elades. he great
llie at
1
New
degree of moiphological di\ersilv in the Zealand hehi-s contrasts with a c()nes[)ondiug lo\s le\el of se<|uence
ilivcrgcui'e. New Ze'aland was a source of new emigrants to t)thcr regions in the South l^acilic that were [)readaplcd to
high mountains or forest margins. Our results suggest that two instances long-distance dispersal from New Zealand
t)f
to South America, at least one instance from New Zealand to Australia, and one instance from New Zealand to New
(iuinea have occurred relati\el\ rec^Mith. Short(M" hops (Chatham Ishmds and the suhantaretie islands arc alst)
tlu^
Itt
supported h) s(Miuence data.
tlie
New
Key words: /A7>e. ITS, Z<'aland, ph\logcuctIc analysis, rbA., Scrophulariaceae, Vewnita.
New
L(nig-dislanc(^ tlispersal has a profound influence in Zealand are conspicuous elein(*nts in most
on evolution of insular floras (Carl([uisl, 1971), terrestrial ecosystems except (orchis and wcllatuls.
llie
and there substantial evidence suggesting that SptM'ies such as UeJw (irmstrongii, H.
is r///»re.s.vo/V/e.s-,
il
occurs ndalively frequently (Godley, 1967; and IL speciosa have patchy or localized dislri-
l\)lc,
One
1991). of the most rcrnarkahle examples of dis- hutions and are c<»nsidered rare or endangered;
persal folloAved by atlaptivc evolution on islands is about 7()7c of llie species are confimnl to small
New New
the Zealand hebi's (Scrophulariaceae). Wags- regions within Zealand,
Carnock-Jones (1098, 2000) suggesttnl The New Zealand hebes were formerly included
laff an<l that
New
the Zealand hcbcs arc the descendants of a in a broadl) dclincd circumscription ol the genus
may Cheeseman,
small founder population have been de- Veronica (Weltstein, 1891; 1925), but
that
&
rived from a single seed. They proposed that com- recent flora and taxonomic treattnenis (Ashwin
biiu'tl influences of irdueeding, genetic drift, and Moore in Allan, 1901; Curnock-Jones, 1993a, b;
strong s(dection acting upon small populali(»ns have Ih^uls, 1994a, b) recognize less in(dusive groups
New
probably played a major role in the diversi- (see Tal)le usually accepting four genera in
i^a|)kl 1),
and
fieation of grouj). Zealand: Cliiofioht'he, Hcbt\ Heliohehe, Ptira-
tin'
Heads
The hebes arc one of largest and most eco- hcbc (Carnock-Jones, 1993a, b). (1987) de-
th<"
New
logically diverse plant grou[is in Zealand, in- scribed an additional genus. Leonohcbe. Altliough
cluding over 120 species, with outlier populations we do not acct^)! his wide circiunserij)tion of that
New name
eastern Australia, Tasnuuna, (/uinea. Ha[)a genus, the Ijconohchc could be aj^plied to a
In
m
Islanil, and South America. I'hev range f al- small clade of (our or five species that is su])pt>rlcd
pine cushion-forming 5D) lowland by the analyst^s of Wagstaff and Garriock-Jones
(Fig. to
i)latils
woody 5M, and
shrubs or small trees (Fig. R, S), (1998. 2000).
Tlie authors gralefull) a<kno\viedge curators of the e\pfM*inicnta1 <j;anlens and ClIK llcrhariuni at Larulcare
*
DNA
Research, Lincoln, along with the assistance of Anita Thome, Atuly (^lazier, and Klizahelh \hicAvoy witli iIk^
se<[uencirig. Dick Olmsleail conli iliuted se\tMal of the ih(\. seqeuiices inchrdcd in our stud), bill Malcolni (Micro-
Optii's Ltd.) provided man) of the f>Iiotographs. arul Alison Kellow assisted with preparation of figures, harlici" versions
of this inanuscri|)t heuelrlcd great!} from the insij^hlful comments of ('hristitie Bc/ar', Use Br-eitwiesiT, Mark (Ihase,
Murray Dawson, and Mison Kellow. fhis rcscaixh was funded in N(*w Zealand h\ the Foundation for R<'sear"ch, Science
and Technology (contracts {'X)9(A\\ and MNZ6()1|. and the Mars<len Fund (lontract LAN601).
New
^I.andcare Research. P.O. Box 69, bincttln 8152, Zealand. vva^stafr>^landcart'.cri.nz.
Mu-^cuni of New Zealand Tc Papa Ton^^arewa, P.O. Box New Zealand.
1(>7, Wclliri'iiton,
'
Scliool of I^iological Sciences and Island Biolog} Research Progranirne, Victoria TniNersity ol Wellington, P.O. Box
'
New
Wellington. Zealand.
()(M),
W
. 1 Botanisches histilut der rnivcrsitat \\ icn. Ri'rrrrwe"; 14, 1().'?() ien. Austria.
Ann. MissoLKi Bor. Gamp. 89: 38-63. 2002.
Volume Number
Wagstaff 39
89, et
1 al.
New
2002 Hebes
Zealand
Diversification of the
New The New
Tal>lt^ 1. ClassifitaUons of Zealand liebes. Zealaiul species were placed in three sections of \^c'ltstein's
(1H91) treatment of Veronica, and Cheeseinan (1925) placed thein In two divisions of Veronica,
&
Ashwin Moore
Weltstein (1891) C^heeseman (1925) in Allan (1961)' Heads (1987, 1994b) Garnock-Jones (1993a, b)
Veronica Veronica
sect. Pygmea Division Pvi^mea Pygtnea Chionohebe Chionohebe ludin^
liu
Parahebe "Group
B")
Hebe Leonohebe
sect.
Denslfoliae
sect.
Hebe
Hebe eb
Division II
<9
Leonohebe
''Seniifla^rih^rmes sect. "Seiniflagrifornies"
Lonnatae Connolae
Lorniatae
sect.
sect. Apili
1^
"Fhigrilormes Flagriformes "Flagriformes •>t
sect.
Aromaticae
sect.
Salicornioides
sect.
i( 9»
**Bu\ilohatae" sect. Baxijoliatae Buxifoiiatae
Hebe
»9
"Subihstichae Subdbtichae
sect. sect. Subdislichae
"Subcarnosae' sec 67aucae sec Gla ucae
t. t.
Hebe Hebe
sect. sect.
"A[)ertae" Hebe Hebe
ser. ser.
"Ocelusae"
sen Oeclusae Oeclusae
ser.
P
araheb e
"Cranchdorae' "Granrhliorae"
Heliohebe
"PaniiHilatae" Paniculatae
sect.
sect. Chamaedrys Division Eureronica Parahebe Parahebe
"Group A, B, C" "Group A, C"
Derwentia
Labiatoides
sect.
Paederota
sect.
Paederoloides
sect.
Pseudolysimachia
sect.
seel. Veroiucaslriun
Omphalospora
sect.
Beccabnuga
setrt.
New
In the Flora of Zealand \olunic (Allan, 1961), M. B. Ashwin prepared the treatment of Parcdiebe^ Pygmea,
'
1
anil tlie infijrmal grouping "Flagriforrnes'' of Hebe, Tlie remainder of the flehe treatment, including the informal synopsis,
was prepared by L. B. Moore.
New
This research contributes to ongoing efforts to ported nionophyletic groups among the Zea-
create a phylogenetic classification of Scrophulari- land hebes, improve
to their classification, infer
aceae. Olmslead and Reeves (1995) and Olmstead their origin, and explore underlying processes of
We
et al. (2001) sliowed that the Scrophulariaceae, as diversification. propose that diversification in
traditionally circumscribed, are not nionophyletic. the group reflects transoceanic dispersal and adap-
Tliey identify clades from a dismembered Scrophu- tive radiation. The liebes have successfully exploit-
lariaceae that could formal recognition. In ed a diversity of ecological niches that were prob-
s.l. itierit
their studies Veronica was nested within a large ably created during the recent uplift and glacialion
New
clade they called the Antirrhinaceat* nam, cons. of the moinitains of Zealand.
prop. (Reveal et 1999). This large clade was
al.,
recognized by Olmstead and Reeves and
(1995) in- AND MehioDS
Ma'FIJUALS
eludes part or all of Bcntham's (1876) tribes Digi-
Our
laleae, Antirrhineae, Chelojieae, and Cratioleae, sampling strategy capitalized on the unique
the small Angelonieae, and the small families characteristics of rbch and ITS sequences. The
lril)e
Callitrichaceae, Clobulariaceae (excluding Selagi- plastid encoded gene rbch has relatively few vari-
naceae), Hi[)puridaceae, and Plantaginaceae. able which allowed sequence comparisons
sites,
The aim of this research to ich'ntify well-sup- among distantly related outgroups, and placement
is
40 Annals the
of
Garden
Missouri Botanical
The
of llif lu'lx's widiiii Scropliulariaceae. It is also use- llu'se genera were ineludeil in our analysis.
was
large nuniher of published rhcL se- Asiatic species Vcronicastrum sihiriruni desig-
ful in that a
(juenees are available (or comparison (see Chase et nated as the oulgroup for th<* analysis of ITS se-
al., 1993; Kiillersjo et al.. 1998: Olnisteuid et al.. qu<Mices.
2UUK
and refertMU-es therein). Finally, Albert et al. Voucher specimens are listed in App*-ndix 1,
(1994) and Bremer and Cuslafsson (1997) suggest- along with collection information, litt^ralure cita-
ed gene ap|)roaches clock-like behav- GenBank
that the rAcl. and (<http://www.ncbi.idm.nili.
tious.
and hence amount
ior in its evolution, the of se- gov>) accession numbers. The c(miplele data sets
could be used estimate
qu<Mice diverg(Mice to are available upon request from the hrst author, and
divergi'uce times. By comparison, the nuclear en- Thm^BASE
deposited (<http://www.
^^,p,,, in
^\^^.y
man\ more
code<l ITS-reglon has variable sites than
The
heri)aria.harvard.edu/lrccbase>). study acces-
which providi^s mon^ informative characters
r/jcL,
niimlxM- S623, and the matrix accession
^j^,^ is
tavonomic
ivsolve relationships loutM- levels
to at m961
and M9()2
.„-,. (rbcL) (ITS),
,^,„^],,.„^
(Baldwin 1995).
et al.,
AM)
lAlKACriON, AMIMIllCAIiON,
1)^A
STl (,llOLi'
l»^
SKQIKNCINC
The rbcL studv group ct)nsisted of 33 species
DNA
Total was extracted from either fresh leaves
New
Zealand
including 12 of the hel)es with
at It^ast
mod-
or leaf fragments dried with silica gel using a
represenlativc from each of the current!) recog-
1
CTAB
method Doyle and
iHcation of the hot of
S species Veronica, and species
nl/.ed gt^nera.
*.f 1
cpDNA
{VMl). The gene rhd. and the
r>^>)'^
Vcmnicastnim. Nineteen sequences were
of rhcl.
rUNA
tJiDNA ITS-reglon [the 3' end of the 18S
ncul\ published luMvin with 13 published
al(»ng
-1
gene; internal transcrilx^l spacer (lTS-1); the
Olmstead
sequi^ncps of Aniirrhinaceae from ci al.
-2
rONA
and McotUuw tahacum was 5.8S gene; internal transcribed spacer
(Solanac(\ie)
(2001),
rDNA
Seven (lTS-2); and the 5' end of the 28S gene] w<M-e
dcsIgnaltHJ as the oulgronp (Lin el al. 1986).
s(»qnences were considered redundant; even though am])lilicd by PCR. Primer sequences and our am-
Ohn-
and sequencing techniques follow
they were not identicab the resolution of missing plification
We
make and Wagstaff and
data could p<»tentially them identical. stead v\ al. (1992) for r/>cL, (ku'-
Excess
ihcrefore excludtnl Dcnvcntia dcrneuliiitia and D. nock-Jones (1998) for the ITS-region.
and Zealand accessions primers and unincorporated nucleotides were
pcrfoHata, the Ne\\ of /ye/>e re-
PCR
cUipdca and saUcifolia, from subscfjuent analy- moved from the prodiuts by spin column cen-
//.
Nineteen 37 species included
ses. o( the in tlu^ rhc\. trifugation (Ql
GKN DNA
analvsis were also included the ITS survey. rhe samples were then
in purified
Inc.).
The ITS stud) grou[) included 78 sequences, 19 labtded with Big Dye terminators (PE Ap|)litMl Bio-
Among
of which were newly ]>ublished. these are systcMUs, The Perkin-Elmer Corp.). Both the forward
S8 Chlonohchc, Dcnrcntia, He- DXA
representativ(^s of
^„J rexcrse strands were sequiMiced by the
Uohche, and Parahehe. including conspecific acees- DNA
Sequencing Con-
Waikato University Facility.
Chiouohehe and from
sions riliolata C. (lensifolia
(.f and assembly was accomplished using
editing
,5^
Pamhehe
Australia ami N(nv Zealand, Uthophila
(Gene Codes
Sequ(^nch(^r 3.0 Corp.).
\(M'sion
New
from Australia, and /* landcuateri from Guin-
ea, riiirty-five speci<'s of Hehe were also iticluded,
Ai.K.NMKlNr
^f''Ql'KN(.K
and among wcvv: one representative
tlu^sc at least
from each of Allan. 1961) informal
!VIo<ni'*s (in
The alignment ITS-region was
secjiKMice U)i die
gnnips; fonuosd from Tasmania; henthamii
//. //.
A
(Thompson
by ClustalX ah, 1997).
facilitaltnl et
New
from the Zealand subantarclic islands; bar-
//.
and gap extension pen-
gaj) penalty setting of 7.S a
ken^ chalhamii an<l (liejfenbachii from the
//. (L //.
and
New alty of 6.6 were initially usetl to identify po-
Chatham main
Islands (east of the islands ol
New sequet»ce data: then low-
accessions from both ^'«i*»" ^^^''&' '" *''^'
Z<'aland); of//, elliptica g^'P^
scoruip segments were realigned using a ga[^
Zealand and the Falkland Islands; and H. sdlici-
New Zealand ami penalty setting of 15 and a gap extension |)enalty
from both Cliih^ Psi
folia
of 6.6 with the removing new gaps optii)n turned
if^
These opened and positioned small
groups hebes on. settings
(Miicrged as potential sister of the in
Hong and Albach and gaps. The aligmncnt was inspected and minor
the analysis of (19{>1) (^hase final
(2(101); therefore a total of 19 species rejiresenting revisions were mad(* manually.
Number Wagstaff 41
Volume
89, et al.
1
New
Hebes
Zealand
the
2002 Diversification of
maximum
parsimony
DATA WAI/lSIS and one of the trees is
1,
members
diown Figure Most of the Antirrhin-
in 2.
The analyses were ai-eomplished
phylogeiietic
aceae form a basal grade in our analysis witli Cluo-
PAUF* 1W8).
using version 4.0d65 (SwoCford, Tlie
Parahrhe,
Dcnccniia, ^Hebe, Heliofwhe,' anc
^^^.j^^.j,^^
analyses were conducted using the PAUP'^' settings
100%
Veronica f(»rming a clade that receives jack-
random addition sequence with 100 re{ilicates,
knife supj)ort. V anagaUis-aquatica is sister to a
TBK
and
branch swapping, mulj)ars
in cITect,
largely Australasian clade (987^ jackknife supi)ort)
The were unordered
steepest descent. characters all New
Zealand hebes along with
includes the
that
and weighted equally, and gaps were treated as
and Rela-
Veronica arguta, pcrsica.
Deru'cniid,
1;
niissing data.
tionships within this clade are poorl) rcsol\ed
Support for the inferred clades is given by jack-
(Fig.. 1,2).
and
1996) by the
knife [)ercenlages (Farris et al..
The mean absolute distance and standard devi-
mimber synapomorphies each group. Jack-
of for
Chionoh-
Veronicastrnm silnricum
to llu^
.^^j^^^^ ^^.^^^^^
knife analysis investigates the structure, or phylo-
and
Parahche,
Dcrwcnlia, Heliohebe, Vc-
Ilche,
^,^^^,^
matrix without juMinutation, but
genetic signal, in a
ronica terminals is 29.1 7.7 (see Fig. 2).
here
excludes an assigned fraction of characters,
Veronicaslrum n^ported in the fossil record (Tiff-
is
The jackknife searches were i>crfonned
set to ;?07f jr^o-^ Miocene souh^ 15 million
. ^^^^^ ,,^^ ^^jj
f^.^^^^^
w 1000 replications excluding uniuformative
ith The
y^;^^^ present (mybp). substitution rate
1^^,,^^^^.^
=
maxtrees 10 for each replication; the start-
sites, was
in the Veronicastrnm lineage estimated l)y di-
random
ing tn»es were obtained by addition with ± - ±
viding29.1 7.7/15 1.9 0.5 substiluli()ns for
TBR
one each jackknife replication,
re{)lic-ation for
^j^^ ^^^-^^^^ ^^^^ ^^^.|^ j^^^ ^^^.^^^ distance from the
and nmlpars
branch-swap[)ing, In effect.
node Australasian
terminals the ancestral of the
to
The between setiuence divergence
relati(mship
which Furasian)
species (including persica,
is is
V.
and gene rbch was (hscussed by Albert
time for the
which corresponds an upper Mio-
changes,
to
1J5.8
and Bremer and Cnslafsson (1997) and
et al. (1994) cene divergence estimate of about 9.9 mybp. The
was calculated using the equation:
mean distance from the terminals to the ancestral
dian^es.
^"^^"M" 1^"^^^^ ^.4
—
"'^''^ ''f" ''i*' ^'-
substitution rate patristic .listaMie(l)j))/.u,.nI,er if
which corresponds to a Pliocene divergence esti-
of nucleotides/inferred time since clado<i;enesis.
mybp
mate of about 3.9 (Fig.
2).
The abgned ITS matrix was 695 nucleotides in
Resiits
length with gaps created to account for insertions
The sequences were 1402 tiuclcolides in and deletions, among which 364 sites were con-
rbc\.
27-1428 Among 109 were parsimony-uninformalive, and 222
length ([)ositions in tobacco). the stant,
1402 included in the rbc]. matrix, 103 sites were potentially parsimony-informative. Missing
sil(^s
1
and gaps accounted
were 44 were parsim(Miy-uniuformative, data accounttnl for 1,29c for
invariant;
1
The 5.8S gene
and 155 characters were parsimony-informative. 9,b7c of the ITS data matrix. a s
\\
between
Missitig data accounted for 4.7% the matrix. unifoiinly 165 luicleotides; lTS-1 varieil
(»l
203
5' 175 and 224 nucleotides and ITS-2 between
Most of the missing data fell in a region at the
end of rbch, upstream from the conserved EcoRV and 215 imcleotides. Most of the variation in the
The
used some setjuences ITS region was observed in lTS-1 and ITS-2.
restriction site in cloiiing
and downstream 5.8S gene was more conserved. Conse^rved motifs
(Olmstead ah, 1992), in a region
et
PCR primer by Liu and Shardl (1991) and Ilershkov-
from position 1325, the h)calion of a identilied
used ampUfy some secjucnces. Most changes and Zimmer (1996) were idenlihcd in the ITS-
site to itz
maximum and lTS-2 sequences our survey.
across the |»arsiinony tree 1 in
(calculatetl
and
shown in Fig. 2) occurred in the ihiid codon posi- Thirty-eight insertions deletions (indels)
tion (369); substantially fewer (*hangcs occurred in were inferred in the ITS-1 and lTS-2 spacer regions
and second codon positions. (Table Mostl) these were relatively small, rang-
the (116) (47) 2).
first
27
Changes the third codon position are generally ing from 1 to 3 bp, but two large Iiisertions of
in
svnonvmous, and hejice are more likely to evolve and 44 bp and three deletions of 7 or 8 bji were
manner. also inferred. Most of the indels were uni(|ue to a
in a clock-like
more
The analysis of rbcL sequences reco\<T(Ml 9408 single sample, but 13 were shared by two oi
maximum sometimes uniting groups su|)ported by
parsimony single species,
trees (listributt;d in a
=
sequence Species
island of 529 steps (consistency index 0.52 ex- substitutions in the data alone.
eluding uniuformative characters, retiMition index of Denventia are characterized by a on<'-base de-
=
consensus showti Figure letion. Species o{ lleliohebe are characterized by a
0.73); a strict tree is in
a
42
Annals
the
of
Missouri Botanical Garden
Hebe
elliptica
Hebe
benthamii
69
Hebe C/3
odor
a
Hebe
salicifolia
Hebe
salicomioides
Parahebe
vandewateri
Hebe
macrantha
Hebe
cupressoides
N
Heliohebe
raoulii
Chionohebe
densifolia
Hebe
cheesemanii
1
Parahebe
catarractae
Derwentia
nivea
Veronica arguta
Hebe
formosa
Veronica catenata
Veronica persica
Veronica
anagallis
-aquatic
Veronica
officinalis
purpurea
Digitalis
Plantago
lanceolata
Veronicastrum
sibiricum
100
Callitriche heterophylla
Hippuris
vulgaris
85
Chelone
obliqua
98
Collinsia grandiflora
68 Antirrhinum majus
Globularia
cordifolia
Gratiola
pilosa
88
Amphianthus
pusillus
Bacopa
caroliniana
-
Angelonia pubescens
Nicotiana tabacum
Ki^uic
1. SIiIlI LoiiM'iisus of 91()fl niiiiiiiuil length tret's producfMl In parsinuiny analysis of r/nl. sequences. This
New
tree shovN^ the plaeenuMit of Zealand hebes Anlirrhinaeeae sensn
withiji tlie Olriislead et al. (2(H)I), using A7rn/mrK7
>
taluirum as an onlgroup. Jaekknifo \alues 50% are given ahove eaeli node.
a
43
Volume Number Wagstaff
et
89, al.
1
New Hebes
Zealand
2002 Diversification of the
r
Hebe
benthamii
It
Hebe
salicomioides
Hebe odora
Hebe
elliptica
Hebe
3.9 salicifolia
'
mybp
Derwentia
nivea
Veronica arguta
Parahebe
catarractae
Parahebe
vandewateri
Hebe
cupressoides
Hebe
macrantha
Chionohebe
densifolia
Veronica catenata
Veronica persica
Hebeformosa
Hebe
cheesemanii
9.9
Heliohebe
raoulii
mybp
Veronica anagallis -aquatic
Veronica
officinalis
Veronicastrum sibiricum
15
mybp
purpurea
Digitalis
Plantago
lanceolata
Callitriche heterophylla
Hippuris
vulgaris
Globularia
cordifolia
Chelone obliqua
Collinsia grandiflora
Antirrhinum majus
Gratiola pilosa
Amphianthus
pusillus
Bacopa
caroliniana
Angelonia pubescens
—
tabacum
Nicotiana
changes
5
The
Figure One of the maxiniiim i^arsimony trees reeoveretl from a parsimony analysis o{ rbch sequi^ices. braneli
2.
Veronuastrum
l(Migllis are proportional to llie rmniher of changes along eaeh branch. See scale at bottom. Fossils ol
silnricunt are reporte<l from the mid Miocene about IT) m)bp. Divergence estimates are provicled al the ancestral n(>(l(^
node Hebe
of Australasian species persica is Eurasian) and the ancestral of the clade.
(I.'
44
Annals
of the
Missouri Botanical Garden
Tabic 2. Insertions ami deletions infenvd ITS
(Votn setjnetiee cdiiiparison.
liisptilon/
iMHl
Ti (Jelt'tidn ze Posili on Setjnence
Si/.
m
Veronirastnuu rim
si hi Dt'lflimi 41
1
t/.
Vrmninfstrum
sibirinim, Wulfrnia Insertion 54
1
c/in b ara
'n I i
Veronica ghuidulosa
67
DcIl'IIoii
1
tvrnicosa 68-79
tr
lli'l
Iii.scition gla
'nmua
al) ciilia Inscrlion 77
c 1
Vvronica (haniardrys
Delclion 8
1 r/g
i
\ert)/iii-astnim sihiriatrn 89
Delt'lion
I
^uljvtiia carintlnai'a Insertion 44 90- 34
aalelagptglgraagcccelti^U^a^ag-
1
tueegc^CL'lgctc
Pscitdolysimaihion
spicafa, Irronica 27 08
Itisoitioii 3 gaela^N-^agt^e^eegeli^c'gcle
1 1
I
anagallis-aquatica,
Iwllidioides,
\. V,
glandulosa,
glaiira,
V. V, offuinalis,
V.
srrpyllifolia,
V, urlicifolia
Vrnnu'cft hellidioidcs, offuinalis, Insertion 162
V. V. 1 a
urticijoiui
Dementia
deruefitiana, niira, D. Insertion 207
I), 1 a
i^inthehe
perfoliata, Uthopliilii,
Veronica arguta
Veronica
tnacrostacli \a
Deletion 3 20JI-209
cc
Hehe
salicifolia Insertion 217
1
Vemniea
bellidioides, glandnlosa, 231
Insertion
1. 1 c/g
V V
officinalis, nrtieifolia, Wulfenia
earinthiaca
Veronica persica Deletion 248
1 a
Veronica chaniaedrys, anagallis- Insertion 260
\'. 1 a
aijuaUca
Psendolysimailiion
spicaia, Veronica Insertion 442
1 a
V
chaniaedrys^ macrosiachva. K
of ens
is
I
\eronica
nrtieifolia Insertion 454
1 e
Ven)nica
anagallis-(t(jnatiea
Insertion 167
1
t
Pseudtilysiniaehion spirata, \eroniea 474
)eletioii
1
I
V
anagallis-aquafiea.
hellidioides,
V.
frntienlosa, glandnlosa, glanca,
V, V.
V
V
njjieinalis, safnrejoides,
V.
serpyllijolia,
nrtieifolia,
\[
Veronieastrum
sihiricnni, Wulfenia
earinthiaca
Parahehe
canescens
Insertion 475
1 (T
eronica
hellidioides
Insertion 2 501
I re
Wnlfcfua earinthiaca
Deletion
1 5( 13
t
IJelioliebe hnlheana. laraudiana. H. Insertion 508
//.
1
t
ndii
ra<
Veronica persica
Deletion 573
1
t
Uehv
Parahehe
elliptica, elliptica, Deletion 8 583-590
II.
hirleyi, I] hreristylis, P. decora, P. lyallii,
spathnlata, vandewateri
P. P.
Parahehe
planopetiotata 590-596
Deletion 7
catelcc
Parahehe
canescens
Deletion 8 591-598
aleteei;e
Veronica persica 592-594
Deletion 3
tea
Ih'liohehe hulkeana, laraudiana.
Deletion
11. //. 1 16
( )(
raoulii
Parahehe
landeualerl
Deletion 609
a
1
Va
eroni ch aniae(
lry> Deletion 2 )9-6
\
6(
1
^^i^
45
Volume Number Wagstaff
et
89, al.
1
New
Hebes
Zealand
2002 Diversification of the
Table Continued.
2.
Insertion/
Sequence
Tax on deletion Sl/e Position
615-616
Parahebe canesccns Insertion 3 cat
620-621
chamaedrys Deletion 2
Veronica tc
625-626
Pseudolysimachion spicata Deletion 2 ac
631
Veronica chamaedrys, serpyUifolia Insertion 1 a
V,
638-639
Insertion 2
Veronica austriaca, olteasis
V.
640
Veronica arguta Insertion 1
one-base insertion and a one-base deletion. Both first is a clade that comprises Leonohebe s. str. in-
accessions oi Hebe elliptica and nine species of eluding a well-supp<n'ted group, //e/>e /e/raA7feAa, //.
Parahebe have an eight-base deletion that appears cheesemanii, and H. ciliolata (99% jaekknife; 6
have evolved independently least three times synapomorphies) with Hebe cupressoides weakly
to at
(61%
(Table This deletion lacking in Parahebe lin- supported as their sister jaekknife; 3 syiia-
2). is
A
and catarractae subsp. catarractae and pomorphies). The Chionohebe clade (98% jack-
in
ifolia P.
The American 5 synapomorphies) includes Parahebe plan-
subspecies martinii. South accession knife;
Hebe has a unique one-base insertion opetiolata and the cushion-forming species of
of salicifolia
New
that lacking in the accession of Hebe salicifolia Chionohebe with both the Zealand and Aus-
is
New The Chionohebe
from Zealand. tralian accessions of C, ciliolata.
The ITS sequences our study were evolving B clade consists of Parahebe and both the
in at trifida
New
a faster rate than r6cL. The average rate of c^hange Zealand and Australian accessions of Chio-
per variable site for rbch was 1.7 (tree length of nohebe clensifolia (91% jaekknife; 4 synapomor-
+
532/number 144 The The fourth clade includes 6 species of Par-
of variable sites 155). phies).
(93% synapomorphies) and
aligned ITS sequences were shorter than rbcL, ahebe jaekknife; 7
&
there were more variable sites (331), and the av- accommodates the informal "Groups A C" of
erage rate of change per variable site was 3.6 fur Ashwin (in Allan, 1961; Table 1) and P. spathulala.
the ITS region. The fifth clade includes all the species of Heliohebe
(100% synapomor-
Parsimony analysis of the ITS-region recovered in our analysis jaekknife; 13
6931 maximum parsimony trees distributed in at phies). The sixth includes the remaining species of
New
two islands of 1213 steps (consistency index Hebe with l)oth Zealand and South American
least
—
Hebe and Hebe
0.41 excluding uninformative characters, reten- accessions of salicifolia elliplica
— consensus shown (100% jaekknife; 11 synapomor{)hies). Hebe ma-
tion index 0.73); a strict tree is
maximum
in Figure 3 and one of the parsimony crantha is weakly supported as the sister to the rest
(50% 4 synapomorphies)
Figure Based upon the results from anal- of this clade jaekknife;
trees in 4.
ysis of rbcL sequences (Figs. 1, 2), Veronicastrum (Figs. 3, 4).
The
sibiricum was designated as the outgroup. ear-
divergence within the ingrouj) between Wid-
liest is DISCUSSION
fenia earinthiaca and other taxa. fhe Northern
all
Hemisphere species of Verornca are found in hve Large, unwieldy genera with a cosmopolitan dis-
among
clades that form a grade basal to a Southern Hemi- tribution such as Veronica pose the most dil-
sphere clade comprising the New Zealand hebes ficult taxonomic problems for plant systematisls,
One
and A heterogeneous Australian whose opinions are often strongly held. of the
th(Mr relatives.
most vexing of these problems the inconsistent
is
f<
ahebe lithophila, Veronica arguta. and V calycina means by whic:h taxonomists defme generic bound-
(82% jaekknife; 18 synapomorphies) sister to the aries and die recognition of rank within a hierar-
is
New scheme. Recent taxonomic
Zealand hebes, though there relatively chical classification
is little
Hemisphere have
support relationship (70% jaekknife; 10 treatments in the Southern fa-
for this
and
vored narrow circumscri{)tions, several tiew
synapomorphies)
(Figs. 3, 4).
among genera have been segregated from Veronica {see Ta-
Six well-supported clades are identified
New whereas taxonomists Europe and North
the Zealand hebes, but the relationships ble in
1),
amon these clades are unclear (Figs. 3, 4). The America have traditionally embraced a broad ge-
a
46
Annals
of the
Missouri Botanical Garden
corrhimii
fh'hi'
Hche
salic ijolia
Ucbe
salu ijoim
Ht'hc parvijlora
speciosa
W'b<'
Hi'bi' piniL'U't)i(It'x
lownsonti
Hi'hi'
Ht'hc odora
Hcht' iiuiinvtufsa
p
he
Hi' 'eepacridea
Hi'he
rgftuf.fissima
Hebe
c Hebe
HHeebbee
e^lnl.i^p/fnica
Hebi'
dieffen biiihii
fuitha
tii'ht'
(
Hebe
co( kiixneana
Hebe
barkeri
ebe saliconui)ides
ebe artnuhita
ebe annsinmiiii
liebe vemifosit
Hebe
propmqitq
Hebe ^
necfoni
tlehe hcopodiqides
Hebe
necfoni
-'
Hebe
benlluffnii
Hebe
pefnei
Hebe macninlba
Hebe mac ram
hit
Helijfhebe bulkeanu
Heliohebe
Heliphebe iavamiiana
Heitohebe
raaulii
Varahebe
spaliiidaia
Piirahebe
spatbidiita
Parahebe hookeriima
Vcinihehe caiaiructae ^
Pafahebe ora
de{
Parahebe
Parahebe
hallii
.
Parahebe
catarractae
'
Parahebe
cafiescois
Parahebe
Itnijolia
Parahebe
brevisfylis
Parahebe
vandeVvaferi
]*arahebe heesemanii
i
CJupnohehe
densijoiia
B
Chionohebe
'hianohebe
C den\ifi>{ia
J\uahebe
trifida
Partihebe
birlc\i
Chipnohebe
thonusc/nii
Chi/mohebe
pulvinaris
A
Chionohebe
Chionohebe
rUiolata
Chionohehe
ciliolaia
Parahebe
planopetiolafa
Hebe
tetra.slirhu
Hebe
ciliohita
Leonohebe
Hebe heesenman a
(
Hebe
cupressso ides
am
Veronica vufa
Veronica cdtycina
Denveniia
denvcnlia/ia
enxenlia
petfi>liala
\n\entia nivea
\irahebe
lilhophila
Hebe
forwosa
Veronica austriaca
Verotiisa mac hya
ro^siai
veronna
tdlensis
X/'enmica persua
Ver(mica chwnaedn.s'
Veronica
frutiiuiosii
Veronica safurejoides
\\eudoly\imacnion
ala
I \pii
Venmii'a vfaina
Veronica neUidioides
Veron
ica ofju ipalis
Verrfnic a articifojia
Veron ica ^lanaulosa
Venmica
ivtaiiallis (iKiuati
(
Xertmica
\erp\lliJolta
WiJjenia carihrhiaca
Venmicastrum
sibiru urn
6931
Fipiin- W. SiricI roiisLiisus of iiiiiumal U'liglli {vvvs pnKliic«'<l [>arsinionv aruilvsis of entirr ITS-region.
1)\ llie
>
Notable
cladi's are uk'ntifu'd with hrackcls. Jackkiiifc \alues SO'/r are given ahove eaeli node. ^Pnrnhcln' catar-
racfdc suhs[>. nxirlinii. '-Pardhrhc rofarrarfae suhsp. catarractae, 'Hchc hcctorii sul)sp, suhsiniilis, ^llcbe licctoru
siil>^p. hrct(>rii.
ncric defmilioti of Icronira, e.p., W'ctblein (1891). well as Eurasiiui genera Paedcrotd and Pseu-
llie
Tliis (liserepaiiey of opinion eonlrihules lo laxo- dolysimarhinn, and the North American genera
notnie and and One
ar]i!)iguity inslahility. Synlliyris Bcsseya.
[)ossil)le soluli(»n [o
\^
Our
results support those of Alhaeli and Chase lump them broad
all in a eii-eumseription of Ve-
implying genus
(2001), that llie Vrronica is at l)(^st ronira. This move, liowever, would er(vite a eas-
paraph)letie h) exclusion of the Southern Ilemi- cade of nomeuc latural (dian g(*s requn-nig ine rec-
ll
sphere genera Dementia, many
Ch'n)tiohch(\ flehe^ Ile- ognition orf n(^w^ combinations and the
and We
linhehe, fj'onohehe, Ptuiihehe (Figs. as ado[)tion of old combinations within Vemniau
1, 3).
47
Volume Number Wagstaff
89, et al.
1
New
Hebes
Zealand
2002 Diversification of the
Hche
corrigann
Hebe Chile
saiicifolia
Hebe
saiicifolia
pannjlora
'cbe
'
ebe speciosa
L Hebe albuans
Hebe
piweleoides
Hebe
*- townsonii
Hebe cockaxneana
Hebe
J odor'a
Hebe pauciramosa
P-
Falkland
Hvhe
L_r" epacrid(,'a
Hebe ramosissima
*
Hebe
ellipfica Islands
Hebe
{diiptica
Hebe Chatham
diefjenbachii
P
V— Hebe (halhqmica
Hebe
barkeri
»-
Islands
Hebe propinqua
Hebe
hec (orii 4.
Hebe lycooppooadioides
Hebe hedorii ^
Hebe
verrucosa
1/2
Hebe
salicornioides
r
Hebe
annulata
-(-
Hebe armslnmgii
Subantarctic
I
—
Hebe beplhanui
0^
Hebe
petnei
Islands
Hebe macranlba
BHebe
Diacranthii
Heliohehe hulkeuna
7^
Heiufhebe iavauaiana
Heliobebe
riufulii
Parahebe
spafhidata
-2
Parahebe
spafhulata
Parabebe hookericwa
2
Parahebe calarradae
Parahebe decora
/^arahebe
haJlii
SI
%
arahebe
c'atarractae '
Parahebe canescens
Parahebe
hnifolia
Parahebe
brevi.s/xlis
New
Parahebe vandewateri Guinea
.
Parahebe cheesewanii
Chionohehe Australia
densifolia
1
pJ* Chionohehe
deiisijolia
*— Parahebe
tnfida
Pafahebe
' birleyl
Chionohehe ihomsonii
Chwnohebe
pulvinaris
Australia
Chiotiohehe cdiolata
Fi
—\ L Chionohehe cdiolata
Parahebe planopeliolaia
Hebe
(elrasticha
Hehe
cdiolata
Hebe
chee.semanii
Hebe
cupressoides
—
Veronica arguta
^^^^^—
{
Veronica cahr ina
Denveniia denxentimta
Australia
Denventia
perfoliata
Derwentia niyea
Parahebe
lilhophila
Hebe
ionnosa
—
yeronlca ausfriaca
—
I
Veronu
— a nuicnhsiachya
J I
Veronica oltensis
I
Veronii a persic a
Veronica chamm*aedrys
r Veronica fruli(idOSa
'^ Veronica salurecjiooiiad^e's
Pseu aolysinuiclnon spu ata
Eurasia
Verpnici\ glauca
Veronica heujcuoiaes
Veronic a officinalis
Vercniica urlicifoha
Veronica glandulosa
Veronica anagallisqaiiatica
{ Veronica serpylhjolia
Wulfenia carinthiaca
Veronicastrum sihiricum
changes
5
New
Figure 4. One of the maximum parsimony trees reeovered from a parsimony analysis of the ITS-region. Tlie
Zealand hebes eom{)rise a well-supported monophyletie group with outliers on the offshore islands and in Australia.
New Guinea, and South Ameriea. Dispersal away from the main islands of New^ Zealand inferred in nine spceies.
is
Rraneh lengths are [iroportional to the number of changes along each ])ranch. See sealc at bottom. ^Parahebe catarracUw
subsp. martind, -Parahebe catarraclae subsp. calarraciae, '''Hebe hectorii subsp. subsimilis, ^Hebe hectorii subsp. heclorii.
and
accept that retaining a paraphyletic Veronica ob- secjiiences describe patterns of diversiliealiun
New
scures phylogenetic rclati(nisliips; however, an al- in the Zealand hebes.
lernative aj)[)n)ach to recognize smaller, less in-
is
NKW
MAJOM CLADES OF /KALANl)
IIKRES
clusive clades as generic segregates of Veronica.
This approacli was ado|)tcd by Hong (1984). Here A lieterogeneous clade composed of Denicnlia,
DNA
we identify major clades supported by the Hehe forwosa, Parahebe lithophila, Veronica arguta.
