Journal of Biogeography (J. Biogeogr.
) (2006) 33, 1259–1265
ORIGINAL
ARTICLE
1
Department of Systematic and Evolutionary
Botany, Institute of Botany, University of
Vienna, Vienna, Austria, 2Departamento de
Ecologı́a Vegetal (Botánica), Universidad de
Sevilla, Sevilla 41012, Spain, 3Biologiezentrum,
Landesmuseum, Johann-Wilhelm-Klein-
Strasse 73, Linz 4010, Austria, 4Department of
Botany, Faculty of Science, Charles University,
Prague 11636, Czech Republic, 5Faculty of
Biological Sciences, College of Natural Sciences,
Chonbuk National University, Chonju, South
Korea and 6Makino Herbarium, Tokyo
Metropolitan University, Hachioji, Tokyo,
Japan
*Correspondence: Tod F. Stuessy, Department
of Systematic and Evolutionary Botany, Institute
of Botany, University of Vienna, Rennweg 14,
A-1030, Vienna, Austria.
E-mail: tod.stuessy@univie.ac.at
INTRODUCTION
The origin of new species on oceanic islands has been discussed
on many occasions (Carlquist, 1974; Grant, 1996). The most
commonly described process is speciation associated with
ª 2006 The Authors
Journal compilation ª 2006 Blackwell Publishing Ltd
Anagenetic evolution in island plants
Tod F. Stuessy1*, Gerhard Jakubowsky1, Roberto Salguero Gómez2, Martin
Pfosser3, Philipp M. Schlüter3, Tomas Fer4, Byung-Yun Sun5 and Hidetoshi
Kato6
ABSTRACT
Aim Plants in islands have often evolved through adaptive radiation, providing
the classical model of evolution of closely related species each with strikingly
different morphological and ecological features and with low levels of genetic
divergence. We emphasize the importance of an alternative (anagenetic) model of
evolution, whereby a single island endemic evolves from a progenitor and slowly
builds up genetic variation through time.
Location Continental and oceanic islands.
Methods We surveyed 2640 endemic angiosperm species in 13 island systems of
the world, both oceanic and continental, for anagenetic and cladogenetic patterns
of speciation. Genetic data were evaluated from a progenitor and derivative
species pair in Ullung Island, Korea, and Japan.
Results We show that the anagenetic model of evolution is much more
important in oceanic islands than previously believed, accounting for levels of
endemic specific diversity from 7% in the Hawaiian Islands to 88% in Ullung
Island, Korea, with a mean for all islands of 25%. Examination of an
anagenetically derived endemic species in Ullung Island reveals genetic
(amplified fragment length polymorphism) variation equal or nearly equal to
that of its continental progenitor.
Main conclusions We hypothesize that, during anagenetic speciation, initial
founder populations proliferate, and then accumulate genetic variation slowly
through time by mutation and recombination in a relatively uniform
environment, with drift and/or selection yielding genetic and morphological
divergence sufficient for the recognition of new species. Low-elevation islands
with low habitat heterogeneity are highly correlated with high levels of anagenetic
evolution, allowing prediction of levels of the two models of evolution from these
data alone. Both anagenetic and adaptive radiation models of speciation are
needed to explain the observed levels of specific and genetic diversity in oceanic
islands.
Keywords
Amplified fragment length polymorphism, anagenesis, cladogenesis, endemic
plants, genetic differentiation, habitat heterogeneity, island biogeography, island
evolution, speciation.
adaptive radiation (Stuessy & Ono, 1998; Schluter, 2000), in
which from a single introduction several lines of speciation
occur, each driven by selection within markedly different
ecological zones. Morphological divergence among species in
this process is often dramatic, but overall genetic differentiation
www.blackwellpublishing.com/jbi 1259
doi:10.1111/j.1365-2699.2006.01504.x
T. F. Stuessy et al.
may be low (DeJoode & Wendel, 1992; Frankham, 1997;
Baldwin et al., 1998; Crawford & Stuessy, 1998; Emerson,
2002). Well-known examples include the lobelioids and
silverswords in Hawaii (Givnish et al., 1995; Carlquist et al.,
2003).
A contrasting mode of evolution is one in which a founder
population arrives on an island and simply diverges through
time without further specific differentiation. This anagenetic
speciation (also known as simple geographic or phyletic
speciation) has been suggested as an important mode of
evolution in the endemic vascular plants of the Juan Fernandez
Islands (Stuessy et al., 1990), but the pattern has not been
studied in other oceanic archipelagos and is less well under-
stood regarding its genetic consequences than is the classical
model of adaptive radiation. The term anagenetic speciation
has been selected for convenience in communication. It is
realized that the source populations could also be undergoing
change, especially if the environment were to be changing
rapidly, for example as a result of Pleistocene events. If
speciation were to occur in the continental region, therefore,
one might label this simply cladogenesis, involving the
production of two new species, one on the islands and one
on the continent. We do not object to this point of view, but it
is likely that in most cases the ecology of the newly formed
islands will be changing faster and more dramatically than that
of the continental source area, and, therefore, speciation may
be occurring in the islands more often or faster than in the
source region. We recognize these definitional problems, but
they should not obscure the main point of this paper, which is
to focus on what is occurring during speciation on the islands
after colonization. We regard the term anagenetic speciation as
useful to contrast with speciation during adaptive radiation,
which clearly involves the splitting of lineages (or cladistic
divergence) in a more dramatic fashion.
This report surveys the angiosperm floras of 13 island
systems, of both oceanic volcanic (Canary, Cape Verde,
Galapagos, Hawaii, Juan Fernandez, Madeira, Ogasawara, St
Helena, Tristan da Cunha, Ullung) and continental (Chatham,
Falklands, Taiwan) origins, revealing that anagenetic speciation
conservatively explains 25% of the total endemic species
diversity. We seek correlations with physical and environmen-
tal parameters that might explain observed levels of the two
patterns of speciation within each island or archipelago. We
also offer a hypothesis, based on available data, for genetic
consequences of the anagenetic speciation model.
METHODS
A data base of 2640 endemic species for the island survey was
prepared from published floras for 13 island systems plus
additional literature: Canary Islands, Wildpret de la Torre &
Del Arco Aguilar (1987), Coello et al. (1992), Hansen &
Sunding (1993); Cape Verde Islands, Bernard-Griffiths et al.
(1975), Boekschoten & Manuputty (1993), Brochmann et al.
(1997); Chatham Islands, Wills-Johnson et al. (1996); Falkland
Islands, Moore (1968), Lawrence et al. (1999); Galapagos
1260
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Islands, Wiggins & Porter (1971), Hamann (1981), Perry
(1984); Hawaiian Islands, Wagner & Funk (1995), Wagner
et al. (1999); Juan Fernandez Islands, Skottsberg (1953),
Stuessy et al. (1984), Marticorena et al. (1998), Greimler et al.
(2002); Madeira, Press & Short (1994), Hughes & Malmqvist
(2005); Ogasawara Islands, Ono & Okutomi (1985), Takayama
et al. (2005); St. Helena, Cronk (2000); Taiwan, Huang (1994),
Huang et al. (2004); Tristan da Cunha Islands, Wace &
Holdgate (1958), Wace (1961), Miller (1964), Gass (1967),
Groves (1981), McDougall & Ollier (1982), Roux et al. (1992);
Ullung Island, Kim (1988), Sun & Stuessy (1998).
Data on natural vegetation types, ages of islands, and other
environmental parameters were taken from the literature cited
above. Comparing vegetation heterogeneity among archipela-
gos is difficult because different systems of classification of
vegetation are in use. We extracted from the literature the
number of vegetation types of comparable hierarchical level
from each island/archipelago. To reduce problems of compar-
ability we then grouped the number of vegetation types into
classes: class 1 (1–4 vegetation types); 2 (5–8); 3 (9–12); 4 (13–
16); 5 (17–20); and 6 (>20).
The most effective way of knowing whether a species has
evolved anagenetically or cladogenetically is by in-depth
studies of each group, including comparisons with continental
progenitors. Such a detailed level of understanding is not
available comprehensively for any island system. For the
purposes of this study, therefore, we have taken a simple
(conservative) approach to estimating levels of anagenetic
speciation. Single endemic species on an island or archipelago
are assumed to have evolved by anagenetic speciation. This
does not account for possible loss of species as a result of
extinction. This is an important point, as it is likely that Pacific
oceanic (and other) islands have undergone considerable
reduction in both surface area and ecological habitat breadth
during their ontogeny, and specific and genetic diversities have
surely been lost (Stuessy et al., 2005). Hence, some single
endemic species may be merely the surviving member of a
once larger endemic group. Unfortunately, without fossil
evidence, which is rarely available from plants on islands, it is
simply impossible to determine levels of extinction in partic-
ular groups. Two or more endemic species in a genus within an
island or archipelago are regarded as reflecting cladogenesis, if
the assumption is made that they both have descended from
the same continental introduction. This ignores possible inter-
island anagenesis, and therefore our approach doubtless
underestimates this mode. Endemic monotypic genera have
also been treated as reflecting anagenesis, even though it may
be the case that some of these related genera may represent
radiating lines from one original ancestor to the islands.
In addition to the above analysis, we present analyses of
genetic variation in populations of Dystaenia (Apiaceae) from
Ullung Island and Japan based on amplified fragment length
polymorphism (AFLP; Vos et al., 1995) data (Pfosser et al.,
2005). A few technical comments are therefore in order. It is
now well accepted that AFLPs can provide a sensitive measure
of overall genetic variation within and among plant
Journal of Biogeography 33, 1259–1265
 Anagenetic evolution in island plants

populations (e.g. Tremetsberger et al., 2003; Muellner et al.,
2005). Six populations of Dystaenia takesimana (Nakai) Kitag.
(Ullung Island) and six of Dystaenia ibukiensis (Yabe) Kitagawa
(Japan) were sampled, involving a total of 126 individuals.
These were chosen to reflect the geographic spread of
populations within each species. Total genomic DNA was
extracted from silica-dried leaves following the CTAB protocol
(Doyle & Doyle, 1987), with slight modifications, followed by
standard AFLP protocols (Vos et al., 1995; Tremetsberger
et al., 2004). An initial screening of selective primer combi-
nations yielded three that gave clear and reproducible bands:
EcoRI-ACT/MseI-CAC, EcoRI-ACC/MseI-CTG, and EcoR-
AGG/MseI-CAT. The fluorescence-labelled selective amplifi-
cation products were separated on a 5% polyacrylamide gel
with an internal size standard on an automated ABI 377
sequencer. The data were imported into Genographer (Ver-
sion 1.6.0, http://hordeum.oscs.montana.edu/genographer) for
scoring of the fragments; the results were exported as a
presence/absence matrix for further analysis. The Shannon
P
index was calculated as HSh ¼ ) (pj ln pj), where pj is the
relative frequency of the jth fragment. Tests for significance of
correlations were calculated with SPSS 8.0 (SPSS, Inc.,
Chicago, IL, USA). Analyses of molecular variance (amovas)
and the coefficient of genetic differentiation (FST) were
calculated with Arlequin 2.0 (Schneider et al., 2000). R
package 4.0 d6 (Casgrain & Legendre, 2001) was used to
perform Mantel tests for each species (10,000 permutations),
comparing the genetic matrix of inter-individual Jaccard
distances with a matrix of geographic distances between
individuals in kilometres.
RESULTS AND DISCUSSION
Table 1 shows levels of anagenetic vs. cladogenetic speciation
in 2640 endemic species of 10 oceanic islands and archipelagos
and three continental islands taken from published floras. The
highest level of anagenetic speciation is for Ullung Island,
Korea, at 88%. At the other extreme are the Hawaiian Islands,
at 7%. The average value for oceanic archipelagos is 22%, and
for continental islands it is 31%. Overall, anagenetic speciation
accounts for approximately one-quarter of all endemic plant
species on these islands.
It is important to emphasize that these values may
underestimate the level of anagenetic speciation in some
archipelagos. For example, the most detailed assessment of
cladogenetic and anagenetic speciation, based on more than
25 years of evolutionary studies on many island taxa, is for the
endemic flora of the Juan Fernandez archipelago (Stuessy et al.,
1990). In this study, anagenesis was estimated as 71%, which is
much higher than the figure given here (36%, Table 1), and
presumably closer to the actual percentage. Another island
where it is possible to give an estimate of anagenesis based on
evolutionary studies is Ullung Island, where the difference is
less. The figure given here is 88% (Table 1), whereas our
previous estimate based on evolutionary studies was 100% (Sun
& Stuessy, 1998; Pfosser et al., 2002). For other archipelagos
with larger endemic floras, however, no detailed summaries are
currently possible. The other archipelago that has received
much recent, especially molecular, attention is the Canary
Islands. Even there, however, only a portion of the endemic taxa
has been examined with modern techniques (Santos, 1998), and
the genetic data are therefore insufficient for a complete
summary of modes of speciation.
The different levels of anagenetic speciation in different
island systems allow a search for correlations with physical
environmental factors (Table 1). No significant correlations
exist with the number of islands in archipelagos, their size, age,
or distance from major source areas, these being traditional
factors used to explain total species diversity (i.e. endemic and
native taxa) on oceanic islands (MacArthur & Wilson, 1967).
Likewise, no significant correlation exists with latitude (not
included in Table 1). Only two factors, elevation and habitat

Table 1 Features of island systems, numbers of endemic species, and levels of anagenetic vs. cladogenetic speciation

Number of Size Distance from Age Elevation Vegetation No. endemic Anagenetic Cladogenetic
Island system islands (km2) mainland (km) (Myr) (m) heterogeneity species speciation (%) speciation (%)

Oceanic
Hawaii (H) 8 16,885 3660 5 4250 6 828 7 93
Canary (C) 7 7601 100 21 3710 6 429 16 84
Tristan da Cuhna (T) 4 208 2580 18 2060 3 27 33 67
Juan Fernandez (J) 3 100 600 4 1319 5 97 36 64
Cape Verde (CV) 12 4033 570 10 2829 2 68 37 63
Galapagos (G) 16 7847 930 5 1707 4 133 43 57
Madeira (M) 3 792 630 14 1862 3 96 48 52
Ogasawara (O) 12 99 800 Tertiary 916 3 118 53 47
St. Helena (S) 1 123 1850 15 826 2 36 53 47
Ullung (U) 1 73 130 2 984 2 33 88 12
Continental
Taiwan (TW) 1 35,800 130 5 3950 3 724 29 71
Chatham (CH) 1 963 668 80 294 1 37 62 38
Falkland (F) 2 8500 410 Tertiary 705 2 14 71 29

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T. F. Stuessy et al.

(a) 100 40
Dystaenia takesimana Dystaenia ibukiensis
U

80 30

Shannon index
F
Anagenesis (%)

CH
60 20
OS
M
G
40
J CV 10
T
TW

20
C 0
H
Population 1 2 3 4 5 6 7 8 9 10 11 12

0 Sample size 10 10 10 11 11 10 15 10 10 9 10 10
0 1000 2000 3000 4000 5000
Estimated c. 100 >500 >500 c. 100 <50 <50
Elevation (m)
population size >500 >1000 c. 100 <50 <50 c. 1000

(b)100 Figure 2 Genetic (Shannon) diversity among populations of

Dystaenia takesimana (1–6) and Dystaenia ibukiensis (7–12) cal-
U P
80 culated as HSh ¼ ) (pj ln pj), where pj is the relative frequency
F
of the jth fragment. Data from Pfosser et al. (in press).
Anagenesis (%)

CH
60
S O D. ibukiensis (scattered throughout Japan) fails to show a
M
G
reduction in genetic variation (Pfosser et al., 2005) within the
40 CV J island endemic (number and per cent of polymorphic
T
TW fragments, 117.33, 96.17% vs. 103.67, 86.73%, and Shannon
20
C
index 33.13 vs. 29.13, respectively). Furthermore, the island
populations all behave as one large island population with no
H

0 geographic partitioning of genetic variation (FST ¼ 0.014;

0 1 2 3
Habitat heterogeneity
Figure 1 Correlation of anagenetic speciation (%) in endemic
angiosperms of island systems with (a) highest elevation and (b)
habitat (vegetation) heterogeneity. For acronyms see Table 1.
(vegetation) heterogeneity, correlate significantly with this
pattern (Fig. 1a, Pearson two-sided, r ¼ 0.84, P < 0.01;
Fig. 1b, Pearson two-sided, r ¼ 0.77, P < 0.01). Higher islands
generally have greater habitat diversity, and this stimulates
cladogenesis and adaptive radiation (e.g. in Hawaii). Lower
islands with more uniform environments yield higher levels of
anagenetic speciation (e.g. in Ullung Island). Hobohm (2000)
has shown a positive correlation of species diversity in the
Macaronesian Islands with elevation and number of vegetation
zones.
The genetic consequences of anagenetic speciation are also
of interest. In the model of adaptive radiation, the original
immigrant gene pool becomes divided as a result of ecological
isolation, and the resultant genetic variation within and among
populations is low (Johnson et al., 2000). Even if a founder-
flush model is advocated (Rundle et al., 1998), the resultant
genetic variation within endemic lineages stays reduced.
Genetic AFLP comparisons of a progenitor-derivative species
pair between Japan and Ullung Island, which shows the highest
level of anagenesis (88%; Table 1), suggest another pattern.
A comparison (Fig. 2) between six populations of D. takesi-
mana (Apiaceae; endemic to Ullung Island) and six of
4 5 6 Mantel test with genetic and geographic distances,
RM ¼ 0.042, P ¼ 0.15). Based on the AFLP data, each of the
two populational systems is clearly monophyletic, excluding
multiple introductions for the island endemic. This generic
system was selected because the two taxa are morphologically
distinct (Sun et al., 1997), they have the same breeding systems
(outcrossing, Pfosser et al., 2005), they are the only two species
in the genus, hence eliminating complicating factors such as
hybridization among congeners, and a deletion in the trnL-F
intron and spacer regions in the island taxon strongly suggests
that it was derived from the Japanese species, rather than vice
versa (Pfosser et al., 2005).
Examples of single progenitor-derivative species pairs that
do not fit this genetic pattern, based on allozyme data, include
(1) Rhaphithamnus (Verbenaceae), whereby the Chilean con-
tinental R. spinosus has much more allozymic variation than
the endemic R. venustus of Masatierra Island in the Juan
Fernandez archipelago (Crawford et al., 1993); and (2)
Gossypium (Malvaceae), whereby the Galapagos endemic
G. klotzschianum has less allozymic variation than the
progenitor G. davidsonii from Baja California (Wendel &
Percival, 1990). In these cases, however, it may be that the
reduction of population number and size over several million
years (Cox, 1983; Stuessy et al., 1984) as a result of natural and
human impacts has led to the pattern of reduced genetic
variation now seen (Stuessy et al., 2005). There is also a recent
report (Woo et al., 2002) of reduced allozymic variation in the
Ullung Island endemic Hepatica maxima in comparison with
its Korean peninsula congener H. asiatica. We have examined

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ª 2006 The Authors. Journal compilation ª 2006 Blackwell Publishing Ltd

this species pair with DNA sequences and AFLP data, however
(Pfosser et al., in press, unpubl. data), and see a similarly high
level of genetic variation in the island endemic to that shown
in Dystaenia. More work on H. maxima and its progenitor will
be needed to resolve this discrepancy.
The equal or higher level of genetic variation seen in
species derived through anagenetic speciation suggests a
mechanism for their origin. After an initial founder event,
the established immigrant population will have greatly
reduced genetic variation. In a favourable uniform environ-
ment, populations will proliferate. Over generations, genetic
variation will accumulate through mutation and recombi-
nation (Lande, 1992) in the isolated island populations as a
result of changes in allelic frequencies through drift and/or
selection (if the island habitat is significantly different from
that of the continent). Lack of eco-geographic partitioning
of genetic variation within the island keeps genetic levels
high. The result is eventually a new species, divergent in
genetic and morphological composition from its progenitor,
but harbouring equal or nearly equal levels of genetic
variation.
ACKNOWLEDGEMENTS
This work was supported by grant P14825-B03 from the
Austrian National Science Foundation (FWF) to T.F.S. and by
the Korean Science and Engineering Fund (KOSEF) to B.-Y.S.
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BIOSKETCH
Tod Stuessy is Professor and Head of the Department of
Systematic and Evolutionary Botany, and Director of the
Botanical Garden, Institute of Botany, University of Vienna.
His general research interests are in island biology, evolu-
tion of the flora of southern South America, principles and
methods of classification, and systematics of Asteraceae. His
recent research focuses on patterns of genetic variation
within the Juan Fernandez (Robinson Crusoe) archipelago,
phylogeny and character evolution within the ancient
complex of Asteraceae (Barnadesioideae), and phylogenetic,
populational and biogeographic studies in Hypochaeris
(Asteraceae, Lactuceae) of Andean and southern South
America.
Editor: José Marı́a Fernández-Palacios
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