Journal of Biogeography (J. Biogeogr.) (2009) 36, 1084–1099
ORIGINAL
ARTICLE
Welcome back New Zealand: regional
biogeography and Gondwanan origin of
three endemic genera of mite harvestmen
(Arachnida, Opiliones, Cyphophthalmi)
Sarah L. Boyer* and Gonzalo Giribet
Department of Organismic and Evolutionary
Biology and Museum of Comparative Zoology,
Harvard University, Cambridge, MA, USA
ABSTRACT
Aim Biogeographers have long been intrigued by New Zealand’s biota due to its
unique combination of typical ‘continental’ and ‘island’ characteristics. The New
Zealand plateau rifted from the former supercontinent Gondwana c. 80 Ma, and
has been isolated from other land masses ever since. Therefore, the flora and
fauna of New Zealand include lineages that are Gondwanan in origin, but
also include a very large number of endemics. In this study, we analyse the
evolutionary relationships of three genera of mite harvestmen (Arachnida,
Opiliones, Cyphophthalmi) endemic to New Zealand, both to each other and to
their temperate Gondwanan relatives found in Australia, Chile, Sri Lanka and
South Africa.
Location New Zealand (North Island, South Island and Stewart Island).
Methods A total of 94 specimens of the family Pettalidae in the suborder
Cyphophthalmi were studied, representing 31 species and subspecies belonging to
three endemic genera from New Zealand (Aoraki, Neopurcellia and Rakaia) plus
six other members of the family from Chile, South Africa, Sri Lanka and
Australia. The phylogeny of these taxa was constructed using morphological and
molecular data from five nuclear and mitochondrial genes (18S rRNA, 28S rRNA,
16S rRNA, cytochrome c oxidase subunit I and histone H3, totalling c. 5 kb),
which were analysed using dynamic as well as static homology under a variety of
optimality criteria.
Results The results showed that each of the three New Zealand cyphophthalmid
genera is monophyletic, and occupies a distinct geographical region within the
archipelago, grossly corresponding to palaeogeographical regions. All three genera
of New Zealand mite harvestmen fall within the family Pettalidae with a classic
temperate Gondwanan distribution, but they do not render any other genera
paraphyletic.
Main conclusions Our study shows that New Zealand’s three genera of mite
harvestmen are unequivocally related to other members of the temperate
Gondwanan family Pettalidae. Monophyly of each genus contradicts the idea of
recent dispersal to New Zealand. Within New Zealand, striking biogeographical
patterns are apparent in this group of short-range endemics, particularly in the
South Island. These patterns are interpreted in the light of New Zealand’s
turbulent geological history and present-day patterns of forest cover.
*Correspondence: Sarah L. Boyer, Department
of Biology, Macalester College, 1600 Grand
Avenue, Saint Paul, MN 55105, USA.
E-mail: [email protected]
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Keywords
Aoraki, Cyphophthalmi, Gondwanan biogeography, Neopurcellia, New Zealand,
Opiliones, Pettalidae, phylogeny, Rakaia, systematics, vicariance.
www.blackwellpublishing.com/jbi
doi:10.1111/j.1365-2699.2009.02092.x
ª 2009 Blackwell Publishing Ltd
Biogeography of New Zealand’s mite harvestmen
‘With regard to general problems of biogeography, the biota of New
Zealand has been, perhaps, the most important of any in the world.
It has figured prominently in all discussions of austral biogeography,
and all notable authorities have felt obliged to explain its history:
explain New Zealand and the world falls into place around it.’
(Nelson, 1975, p. 494)
INTRODUCTION
New Zealand has intrigued students of biogeography since the
days of Ernst Dieffenbach (1811–55), the German geologist
and physician who became the first European naturalist to live
and work in New Zealand (1839–41). He corresponded
extensively with Charles Darwin about a wide range of topics,
including the relationship of the avifauna of the Chatham
Islands to that of the New Zealand archipelago. To this day, the
biogeography of New Zealand’s biota remains a subject of
intense interest and debate.
New Zealand’s unique biogeography is a consequence of
the land’s unusual geological history (Gibbs, 2006). Because
New Zealand originated as a fragment of the supercontinent
Gondwana, with rifting occurring some 80 Ma, but has been
isolated from major land masses by some 2000 km for the
past 60 Myr, its flora and fauna have characteristics typical
of both continents and islands (McDowall, 2008). Therefore,
the New Zealand archipelago has been viewed as a unique
system for studying the evolution of a continental fauna in a
state of isolation usually seen only in volcanic islands
formed in the mid-ocean (Daugherty et al., 1993; Cowie &
Holland, 2006). But is there truly a Gondwanan character to
the New Zealand flora and fauna? This topic has been the
subject of fierce debate over the past few years, with some
maintaining that Gondwanan vicariance has played a major
role in establishing the biota of New Zealand (Harvey, 1996;
Cooper et al., 2001; Ericson et al., 2002; Stockler et al.,
2002) and others insisting that submersion of New Zealand
during the Oligocene and/or long-distance dispersal of
organisms to the archipelago have erased any Gondwanan
signal in extant communities (McGlone, 2005; Waters &
Craw, 2006; Knapp et al., 2007; Trewick et al., 2007; Landis
et al., 2008).
Like many isolated islands, New Zealand is characterized by
a biota that is depauperate in certain higher taxa, with rates of
endemism at or near 100% in many groups (Daugherty et al.,
1993; Myers et al., 2000; Gibbs, 2006). Radiations may have
been driven in part by fluctuations in global sea level during
the Cenozoic, as well as the dramatic Oligocene ‘drowning’ or
‘bottleneck’ period when New Zealand’s land area shrank to a
tiny fraction of its present size. An often-cited estimate of the
smallest land area is 18% (Cooper & Millener, 1993), although
recently some have argued that there is no evidence that New
Zealand was not entirely submerged during this time (Landis
et al., 2008). As land re-emerged during the Miocene and the
climate warmed, the diversity of plants and animals increased
(Cooper & Millener, 1993), with radiations in such notable
groups as ratite birds and giant wetas occurring during this
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
time (Cooper & Cooper, 1995; Trewick & Morgan-Richards,
2005). The Pliocene saw the uplift of most of the major
mountains seen in New Zealand today, including the Southern
Alps, and the Pleistocene was a period of cycling glacial and
interglacial stages. Together, these events drove diversification
in many terrestrial invertebrate groups (Trewick et al., 2000;
Morgan-Richards et al., 2001; Trewick, 2001; Trewick &
Wallis, 2001; Buckley & Simon, 2007). Such diversification
should render a clear phylogenetic pattern of reciprocal
monophyly of both the New Zealand clades and their sister
taxa.
The evolutionary history of most plant and animal groups
has been studied with a focus on a single biogeographical
process such as vicariance, dispersal or radiation. One group
with the potential to provide a window into both Gondwanan
vicariance and more recent patterns of diversification is the
family Pettalidae, a group of tiny mite harvestmen (daddy
long-legs in the suborder Cyphophthalmi) (Fig. 1) with a
typical Gondwanan distribution and a remarkably high
diversity in New Zealand. The New Zealand archipelago is
home to 57% of the described pettalid mite harvestman
species, most of which were first documented by local
arachnologist Ray Forster (Forster, 1948, 1952; Boyer &
Giribet, 2003, 2007). Three genera of these animals are
endemic to New Zealand: Neopurcellia Forster, 1948, including
only the widespread species Neopurcellia salmoni, found in the
western South Island between Lake Kaniere and Lake Te Anau;
Rakaia Hirst, 1925, found on Stewart Island, along the south
and east coasts of the South Island and throughout Nelson and
in the North Island as far north as Lake Waikaremoana; and
Aoraki Boyer & Giribet, 2007, with a distribution ranging
throughout the North Island and in the South Island as far
south as Aoraki/Mount Cook in the South Island (Boyer &
Giribet, 2007) (Fig. 2).
Most species of Pettalidae are known from only a handful of
localities, despite extensive collecting by Forster and others,
with ranges typically of the order of 100 km or less in
diameter. This makes these animals short-range endemics as
defined by Harvey (2002). In addition, no species in the entire
suborder Cyphophthalmi are known from oceanic islands
formed de novo by volcanoes, such as Hawaii or the Society
Islands (Giribet, 2000). These slow-moving animals spend
their entire life cycle in forest leaf litter and retreat underground during periods of drought or cold. In New Zealand,
they have been collected from localities close to sea level
through elevations above 1100 m. Although they are not found
in grasslands and shrublands, the authors have collected them
from tiny patches of forest habitat surrounded by agricultural
fields. Therefore, these animals are excellent models for
studying historical biogeography in general, and vicariance in
particular.
In this paper, we present a phylogeny of the family
Pettalidae including 94 specimens, of which 54 are individuals
representing 25 of the 29 known species and subspecies of New
Zealand’s mite harvestmen, plus six undescribed species –
totalling 31 New Zealand taxa. The questions we addressed
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S. L. Boyer and G. Giribet
(a)
(b)
(c)
(e)
Figure 1 Live photographs of selected South
Island specimens belonging to the three New
Zealand genera of Cyphophthalmi. The animals are c. 3–4 mm long. (a) Aoraki healyi
from Mt Stokes (S 4105¢, E 17408¢).
(b) Aoraki longitarsa from Governor’s Bush,
Aoraki/Mt Cook N.P. (S 4344¢, E 17005¢).
(c) Neopurcellia salmoni from Roaring Billy
Falls, Haast River (S 4356¢, E 16917¢).
(d) Rakaia pauli from Kelcey’s Bush,
Waimate (S 4442¢, E 17058¢). (e) Rakaia
stewartiensis from Fern Gully, Oban, Stewart
Island (S 4653¢, E 16806¢). All photographs
by G. Giribet.
(d)
include: (1) How are the New Zealand genera related to other
members of the family Pettalidae from across Temperate
Gondwana, and what can be inferred about dispersal vs.
vicariance within the family? (2) Are the New Zealand
Cyphophthalmi a monophyletic group, or do they constitute
multiple lineages? (3) How can the regional biogeographical
patterns seen within the New Zealand Cyphophthalmi be
related to the geological history of New Zealand?
MATERIALS AND METHODS
Taxon sampling
A total of 94 specimens from the family Pettalidae (suborder
Cyphophthalmi) were included in this study. These included
54 individuals representing 31 species and subspecies of three
endemic genera (Aoraki, Neopurcellia and Rakaia) from New
Zealand, plus other members of the family from Chile (genus
Chileogovea), South Africa (genera Purcellia and Parapurcellia),
Sri Lanka (Pettalus), and Australia (Austropurcellia and Karripurcellia). Members of the families Sironidae, Stylocellidae,
Neogoveidae and Troglosironidae are included as outgroups
(see Appendix S1 in Supporting Information). The majority of
the material was collected alive by the authors and fixed in
96% EtOH, with a few specimens collected by the New Zealand
Department of Conservation (NZ DOC) using pitfall traps. All
specimens are deposited as vouchers at the Harvard Museum
of Comparative Zoology, Department of Invertebrate Zoology
DNA collection, with the exception of the four specimens
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collected by the NZ DOC, which are deposited at the Museum
of New Zealand Te Papa Tongarewa (for voucher numbers see
Appendix S1).
Morphological data set
A total of 45 morphological characters were scored for each
species used in the molecular data set. These characters were
defined in our 2007 study of the phylogeny of Pettalidae and
references therein (Giribet & Boyer, 2002; Giribet, 2003a; de
Bivort & Giribet, 2004; Boyer & Giribet, 2007) (Appendix S2).
When appropriate specimens were available, they were
dissected and mounted for examination using an FEI Quanta
200 scanning electron microscope (SEM). Micrographs are
available in the online public database MorphoBank at http://
morphobank.geongrid.org. Characters were also coded from
specimens examined using light microscopy, including
holotype material where available. The Harvard Museum of
Comparative Zoology online database of photographs of Cyphophthalmi types is available at http://giribet.oeb.harvard.edu/
Cyphophthalmi.
DNA extraction
Total DNA was extracted from whole animals using Qiagen’s
DNeasy tissue kit (Qiagen, Valencia, CA, USA), either by
crushing the individual or one appendage in the lysis buffer, or
by incubating an intact animal or appendage in lysis buffer
overnight, then removing the specimen before proceeding with
Journal of Biogeography 36, 1084–1099
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Biogeography of New Zealand’s mite harvestmen
Figure 2 Distributions of the genera Aoraki
(white with thin borders), Neopurcellia
(white with thick borders), and Rakaia
(black) in New Zealand. The map shows the
collecting localities of specimens from the
current study (circles) as well as museum
specimens (squares) assigned to genera using
morphological diagnoses established by
Boyer & Giribet (2007).
the rest of the manufacturer’s extraction protocol, as described
by Boyer et al. (2005).
PCR and sequencing
Purified genomic DNA was used as a template for polymerase
chain reaction (PCR) amplification of the genes for 18S rRNA,
28S rRNA, 16S rRNA, cytochrome c oxidase subunit I (COI)
and histone H3. The complete 18S rRNA (c. 1.8 kb) was
amplified in three overlapping fragments of c. 900 bp each,
using primer pairs 1F–5R, 3F–18Sbi and 18Sa2.0–9R (Giribet
et al., 1996; Whiting et al., 1997). An additional primer
internal to 1F–5R was used for sequencing, 4R (Giribet et al.,
1996). The first c. 2200 bp of 28S rRNA were amplified using
the primer sets 28SD1F/28Srd1a-28Sb (Whiting et al., 1997;
Park & Ó Foighil, 2000; Edgecombe & Giribet, 2006), 28Sa–
28Srd5b (Whiting et al., 1997; Schwendinger & Giribet, 2005)
Journal of Biogeography 36, 1084–1099
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Aoraki
Neopurcellia
Rakaia
and 28S4.8a–28S7bi (Schwendinger & Giribet, 2005). Sequencing of the 28S rRNA gene was performed with those primers
and some additional internal primers: 28Sa (Whiting et al.,
1997) and 28Srd4b (Edgecombe & Giribet, 2006). 16S rRNA
was amplified and sequenced using the primer pair 16Sa–16Sb
(Xiong & Kocher, 1991; Edgecombe et al., 2002). COI was
amplified and sequenced using the primer pair LCO1490–
HCO2198 (Folmer et al., 1994). The complete coding region
of histone H3 was amplified and sequenced using primer pair
H3aF–H3aR (Colgan et al., 1998). A complete table of primer
sequences can be found in Boyer & Giribet (2007).
PCR reactions (50 lL) included 4 lL of template DNA,
1 lm of each primer, 200 lm of dNTP (Invitrogen, Carlsbad,
CA, USA), 1· PCR buffer containing 1.5 mm MgCl2 (PerkinElmer, Waltham, MA, USA) and 1.25 units of AmpliTaq DNA
polymerase (Perkin-Elmer). The PCR reactions were carried
out using a GeneAmp PCR System 9700 thermal cycler
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S. L. Boyer and G. Giribet
(Applied Biosystems, Carlsbad, CA, USA), and involved an
initial denaturation step (5 min at 95C) following by 35 cycles
including denaturation at 95C for 30 s, annealing (ranging
from 42 to 49C) for 30 s, and extension at 72C for 1 min,
with a final extension step at 72C for 10 min.
The double-stranded PCR products were visualized by
agarose gel electrophoresis (1% agarose), and purified using
Qiagen QIAQuick spin columns. The purified PCR products
were sequenced directly; each sequence reaction contained a
total volume of 10 lL including 2 lL of the PCR product,
irrespective of PCR yield, 1 lm of one of the PCR primer pairs,
1 lL of ABI BigDye 5· sequencing buffer and 0.5 lL of
Applied Biosystems big dye terminator ver. 3.0. The
sequence reactions, performed using the thermal cycler
described above, involved an initial denaturation step for
3 min at 95C, and 25 cycles (95C for 10 s, 50C for 5 s, 60C
for 4 min). The BigDye-labelled PCR products were cleaned
with AGTC Gel Filtration Cartridges or Plates (Edge Biosystem, Gaithersburg, MD, USA). The sequence reaction
products were then analysed using an ABI Prism 3100 or
3730 Genetic Analyzer (Applied Biosystems).
Sequence editing
Chromatograms obtained from the automatic sequencer were
read, and both strands for each overlapping fragment were
assembled using the sequence-editing software sequencher
4.0 (Gene Codes Corporation, Ann Arbor, MI, USA). Sequence
data were edited in macgde 2.2 (Linton, 2005). All new
sequences were deposited in GenBank (Appendix S1).
Phylogenetic analysis
Direct optimization (dynamic homology)
Sequences from ribosomal genes were compared against
secondary structure models, and then split into accordant
fragments using internal primers and some visualized secondary structure features (Giribet, 2002). COI, which showed
length variation, was split into three fragments using areas
where the amino acid sequences allowed us to do so
unambiguously. The protein-coding gene histone H3 showed
no length variation and was treated as pre-aligned.
Data from each gene were analysed in poy ver. 3.0
(Wheeler et al., 2004) using the direct optimization method
with parsimony as the optimality criterion (Wheeler, 1996).
The direct optimization method allows analysis of sequences
of unequal length without a predetermined static alignment.
poy uses a dynamic optimization process that generates
phylogenetic trees by searching for topologies that minimize
total transformation cost under specified parameters (e.g.
indels, transversions and transitions). Rather than performing sequence alignment and tree construction under different criteria and/or models, the same criterion and model
are used consistently throughout the entire phylogenetic
analysis.
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The data for all genes were analysed simultaneously. In
addition, the data for each gene were analysed independently,
and 18S and 28S rRNA were combined in an analysis of
nuclear ribosomal genes. Tree searches were conducted on a
cluster of 30 dual-processor nodes assembled at Harvard
University (darwin.oeb.harvard.edu). Commands for load
balancing of spawned jobs were in effect to optimize
parallelization procedures (–parallel –dpm –dpmacceptratio
1.5 –jobspernode 2). Trees were built through a random
addition sequence procedure followed by a combination of
branch-swapping steps (subtree pruning and regrafting, and
tree bisection and reconnection).
poy facilitates efficient sensitivity analysis (sensu Wheeler,
1995). All data sets (individual genes and combinations) were
analysed under nine parameter sets for a range of indel-totransversion ratios and transversion-to-transition ratios (111,
121, 141, 211, 221, 241, 411, 421, 441). A table describing these
parameter sets can be found in Boyer & Giribet (2007).
Implied alignments (Wheeler, 2003) can easily be generated for
each tree (Giribet, 2005).
To identify the parameter set or ‘model’ that maximized
congruence among all the data (what we call the ‘optimal’
parameter set), we employed a character-congruence technique
which is a modification of the incongruence length difference
(ILD) metric developed by Mickevich & Farris (1981). The
value is calculated for each parameter set by subtracting
the sum of the scores of all partitions from the score of
the combined analysis of all partitions, and normalizing it
for the score of the combined length. This has been interpreted
as a meta-optimality criterion for choosing the parameter set
that best explains all partitions in combination, maximizing
overall congruence and minimizing character conflict among
all the data. This parameter set was given special consideration
in the analysis of data from each individual gene, and is
referred to throughout the paper as the ‘optimal parameter
set’. Although Wheeler (1995) proposed the use of such a
measure as an option for choosing between multiple hypotheses, such a measure has been modified subsequently (e.g.
Wheeler & Hayashi, 1998). The ILD has been compared with
other measures by Aagesen et al. (2005), and criticized for
being partition-dependent (Wheeler, 1996). However, we use
the ILD test as a rough way to choose a tree, yet still consider
the remaining parameter sets by evaluating the alternative trees
obtained under the different parameter sets and by presenting
the strict consensus of all trees obtained under the different
analytical parameter sets (interpretive functions of Wheeler,
2006), as discussed by Giribet (2003b).
We performed an analysis of the morphological data set
alone, as well as a combined analysis of molecular and
morphological data, in which all trees from across parameter
sets used in the analysis of molecular data were used as the
basis for tree fusing under each parameter set, with morphological character transitions weighted identically with the
maximum weight allowed for any molecular transition. As in
the combined molecular analysis, the ILD was calculated and
used to identify an optimal parameter set. Nodal support for
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Biogeography of New Zealand’s mite harvestmen
all topologies was measured using parsimony jackknifing
(Farris et al., 1996) with 100 replicates.
Multiple sequence alignments (static homology)
Even though direct optimization approaches such as the one
employed here are compelling philosophically (De Laet, 2005;
Fleissner et al., 2005; Redelings & Suchard, 2005; Wheeler,
2006; Wheeler & Giribet, 2009), they are not wanting for
detractors (e.g. Kjer, 2004; Simmons, 2004; Kjer et al., 2007;
Ogden & Rosenberg, 2007). In order to evaluate further the
hypotheses derived under direct optimization, we performed a
more traditional two-step phylogenetic analysis of the data by
submitting the sequence loci with length variation to a static
alignment procedure. We generated manual alignments for
18S rRNA, 28S rRNA and COI, excluding areas that were
difficult to align visually. For ribosomal data, initial alignments
were generated by splitting the sequences into fragments as
described for direct optimization, and then modifying the
alignments by eye. Alignment of COI was based on translation
of the DNA sequences. For the static alignment of each data
set, as well as for the entire concatenated data set, we
determined the best-fit model using Modeltest 3.7 (Posada &
Crandall, 1998) under the Akaike information criterion (AIC),
as recommended by Posada & Buckley (2004). The model
chosen was GTR + I + G in each case, except for the 18S
rRNA data partition, for which TrN + I + G was favoured. We
then analysed the data under the recommended model(s) using
mrbayes 3.1.1 (Huelsenbeck & Ronquist, 2001), running
sufficient generations such that the average deviation of split
frequencies reached < 0.01. In each case, four chains were
employed, and the number of generations was either 5,000,000
(18S and 28S rRNA data sets) or 2,000,000 (COI and H3 data
sets). The combined (concatenated) data set was run for
15,000,000 generations, and failed to achieve an average
deviation of split frequencies below 0.154. For the combined
data set, each partition was allowed a separate model. In each
case, burn-in prior to levelling of –log L was discarded. We
also analysed the combined data for 18S rRNA, 28S rRNA,
COI and histone H3 using paup* portable version 4.0b10 for
Unix on the Harvard University Center for Genomic Research
supercomputing cluster (portal.cgr.harvard.edu) under the
single model chosen, using the AIC criterion as implemented
in Modeltest 3.7, GTR + I + G. We calculated bootstrap
support with 100 replicates.
RESULTS
In our analyses we tested the monophyly of each New Zealand
genus of Cyphophthalmi: Aoraki, Neopurcellia and Rakaia
(Fig. 3). In addition, several subgeneric clades were retrieved in
all or most analyses. They are described here to facilitate tree
description in this section.
Within Aoraki, two clades were retrieved consistently and
will be referred to hereafter as Aoraki clade a and Aoraki clade
b. ‘Aoraki clade a’ includes animals from Nelson and North
Journal of Biogeography 36, 1084–1099
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Island that all have a longitudinal carina on the anal
plate: Aoraki calcarobtusa westlandica, A. granulosa and
A. tumidata, as well as animals from throughout the northwest South Island: Aoraki denticulata denticulata,
A. denticulata major and A. longitarsa (Figs 3 & 4). ‘Aoraki
clade b’ includes species from the Marlborough Sounds and
North Island: Aoraki crypta, A. healyi, A. inerma and Aoraki sp.
Mount Stokes.
Within Rakaia, four clades were retrieved consistently and
will be referred to hereafter as Rakaia clades a, b, c and d.
‘Rakaia clade a’ is made up of animals with male tarsus IV
bisegmented or bilobed: R. florensis, R. lindsayi, R. minutissima and R. stewartiensis. ‘Rakaia clade b’ is found throughout
Otago and Southland and includes Rakaia sorenseni sorenseni,
R. sorenseni digitata and Rakaia sp. Beaumont Forest. ‘Rakaia
clade c’ includes animals from Otago and Canterbury, mostly
found along the east coast: Rakaia antipodiana, R. macra,
R. pauli and Rakaia sp. Hinewai. ‘Rakaia clade d’ includes
animals from the Wellington area: Rakaia dorothea, R. magna
australis, R. media, R. cf. uniloca, R. solitaria, Rakaia sp.
Akatarawa, Rakaia sp. Kapiti Island and Rakaia sp. Wi Toko
(Figs 3 & 5).
Molecular data
When all molecular data were combined, a well resolved tree
was retrieved under the optimal parameter set (411) (Table 1;
Fig. 3). Aoraki, Neopurcellia and Rakaia were all found to be
monophyletic, and there was not support for sister-group
relationships among pairs of these three genera. Monophyly of
each genus was stable to parameter variation, appearing in the
strict consensus of all shortest trees from analysis under every
parameter set. Each genus also formed a clade in the Bayesian
analyses, with posterior probability > 95% and was well
supported in the likelihood analyses (Fig. 6).
Within Aoraki, two lineages were found – Aoraki clade a and
Aoraki clade b – within which the Marlborough Sounds species
form a paraphyletic grade with respect to the North Island
species (Fig. 4). Within Rakaia, a nested pattern was seen:
(Rakaia clade a (Rakaia clade b (Rakaia clade c + Rakaia clade
d))) (Fig. 5).
Results from analyses of individual molecular data partitions
are not discussed here, and can be found in Boyer (2007).
Morphological data
Morphological data alone resolved the family Pettalidae as
monophyletic and retrieved the monotypic genus Neopurcellia
but not Rakaia, Aoraki, or any of the subgeneric clades of New
Zealand species found in analyses of molecular data. Outside
of the New Zealand taxa, the pettalid genera Pettalus and
Chileogovea were both retrieved – Purcellia was represented by
a single species. Parapurcellia appears as a paraphyletic grade
sister to all other Pettalidae. Aoraki clade b appears as a
paraphyletic grade sister to Pettalus (Sri Lanka) + Chileogovea
(Chile).
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S. L. Boyer and G. Giribet
Figure 3 Phylogeny of the family Pettalidae. This tree represents the strict consensus of 50 most parsimonious trees of 14,701 weighted
steps obtained in direct optimization analysis of all molecular data under the optimal parameter set (411). Numbers on branches
represent jackknife frequencies, with stars indicating 100% support. The three lineages from New Zealand are highlighted. Maps correspond
to the distribution of each New Zealand genus.
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Journal of Biogeography 36, 1084–1099
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Biogeography of New Zealand’s mite harvestmen
Table 1 Tree length and calculated incongruence length difference (ILD) values for the partitions, including total combined molecular data
(mol), 18S rRNA (18S), 28S rRNA (28S), 16S rRNA (16S), cytochrome c oxidase subunit I (COI), histone H3 (H3), combined molecular + morphological data (comb) and morphology alone (morph), at different parameter set values corresponding to those described by
Boyer & Giribet (2007). Optimal parameter sets are indicated in bold.
111
121
141
211
221
241
411
421
441
mol
18S
28S
16S
COI
H3
comb
morph
ILD mol
ILD comb
22198
17655
30314
12436
20211
35371
14701
24659
44234
448
323
513
230
331
529
238
347
561
4352
3592
6318
3011
5206
9518
4650
8271
15613
6068
5186
9374
3486
6027
11024
4080
7200
13322
9728
7402
12312
4893
7451
12409
4919
7510
12530
1194
799
1187
598
799
1187
597
799
1187
22317
18036
30806
12685
20699
35862
15192
25157
44731
100
200
400
200
200
400
400
400
400
0.0182
0.0207
0.0198
0.0174
0.0195
0.0197
0.0146
0.0214
0.0229
0.0191
0.0296
0.0228
0.021
0.03331
0.0222
0.0203
0.025
0.025
ILD-mol, ILD when molecular data are analysed without morphology; ILD-comb, ILD for combined molecular and morphological data.
a
b
Figure 4 Geographical distributions of
clades within the genus Aoraki. The tree
corresponds to the Aoraki clade from Fig. 3.
Black circles indicate localities represented by
specimens included in the present study;
white circles represent additional known
localities for the genus based on museum
specimens. The letter beside each clade
corresponds to the map illustrating the
distribution of that clade.
Combined molecular and morphological data
The optimal parameter set for combined molecular and
morphological data was 111 (Table 1). The strict consensus
under this parameter set was almost fully resolved, with a
monophyletic Pettalidae, Aoraki, Neopurcellia and Rakaia
(Fig. 7). Within both Aoraki and Rakaia, the subgeneric clades
described in the ‘Molecular data’ section were retrieved once
again. Monophyly of each pettalid genus and of most New
Zealand subgeneric clades received high jackknife support, and
relationships among genera received < 50% jackknife support
in every case.
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
a
b
DISCUSSION
Geological context
New Zealand separated from Gondwana c. 80 Ma. Throughout
the existence of the supercontinent, the land that today forms
New Zealand was adjacent to land that today forms Australia
and Antarctica (Sanmartı́n, 2002). During the Oligocene
drowning event c. 27 Ma, the land area of the North and
South Islands was reduced and then expanded again due to
changing sea levels, dwindling to a tiny fraction of its presentday size (Stevens, 1980; Cooper & Cooper, 1995; Trewick &
1091
S. L. Boyer and G. Giribet
d
c
b
a
a
b
c
Morgan-Richards, 2005), although some maintain that New
Zealand could have been submerged entirely (Waters & Craw,
2006; Landis et al., 2008; McDowall, 2008). Over the past 20–
25 Myr, the Australian and Pacific tectonic plates have slipped
c. 500 km relative to one another, with the Australian plate
moving north. The Rangitata Orogeny during the Cretaceous
resulted in large mountains that eroded and subsided through
the Palaeogene. New Zealand featured only limited topographic relief until the Pliocene, when mountain-building
began in the South Island (5–2 Ma), during which time all of
the major axial mountain ranges were formed.
The development of the Southern Alps provided conditions
for the formation of extensive glaciers, which reached their
maximum during the Pleistocene, c. 18,000 years ago (Stevens,
1980; Suggate, 1990; Trewick, 2001; Winkworth et al., 2005).
During the Last Glacial Maximum (LGM), ice sheets covered
much of the Southern Alps and extended to the lowlands of the
central west coast of the South Island. New Zealand’s small size
and lack of large ice sheets at the glacial maximum allowed
vegetation to react quickly to climatic change, with a
minimum of periglacial effects from decaying ice sheets.
Although Pleistocene pollen records suggest that forest was
uncommon in the South Island during the LGM, new data
from fossil beetles challenge this view. The presence of certain
beetle species indicates that forest habitats have been found in
LGM strata at localities previously thought to have been
1092
d
Figure 5 Geographical distributions of
clades within the genus Rakaia. The tree
corresponds to the Rakaia clade from Fig. 3.
Black circles indicate localities represented by
specimens included in the present study;
white circles represent additional known
localities for the genus based on museum
specimens. The letter beside each clade
corresponds to the map illustrating the
distribution of that clade.
dominated by grasslands during the LGM (Marra, 2008). A
major increase in arboreal pollen at localities across the South
Island occurred starting 10,500 years ago, and over the next
1000 years a large proportion of the land mass became covered
with tall podocarp-hardwood forest (McGlone et al., 1993).
Today, New Zealand’s forests are composed of four broad
floristic components: podocarps, hardwoods, Nothofagus (the
southern beech) and Agathis australis (kauri). New Zealand’s
Cyphophthalmi are known from all of these forest types,
although a survey of museum collection records and the
authors’ own field work experiences indicate that where these
animals are found, they are most abundant in Nothofagus
forests and least abundant in the kauri forests of the
northernmost peninsula of the North Island.
Marine inundation, orogeny and other tectonic activity, and
glaciation are all phenomena that would have contributed
towards the fragmentation of the forests of New Zealand. The
biogeography of New Zealand’s Cyphophthalmi fauna would
certainly have felt significant impacts from all of these
processes.
Overview: biogeographical relationships of New
Zealand Cyphophthalmi
The three New Zealand genera of Cyphophthalmi are
unequivocally members of the temperate Gondwanan family
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
Biogeography of New Zealand’s mite harvestmen
Figure 6 Phylogeny of the family Pettalidae. This tree is based on maximum likelihood analysis of combined data from 18S rRNA,
28S rRNA, histone H3 and COI data sets. The overall likelihood score for this tree was 48816.98077. Numbers on branches represent
bootstrap support.
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
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S. L. Boyer and G. Giribet
Figure 7 Phylogeny of the family Pettalidae. This tree represents the strict consensus of 30 most parsimonious trees of 22,317 weighted
steps obtained in direct optimization analysis of molecular plus morphological data under the optimal parameter set (111). Numbers
on branches represent jackknife frequencies; stars indicate 100% support. The three lineages from New Zealand are highlighted.
1094
Journal of Biogeography 36, 1084–1099
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Biogeography of New Zealand’s mite harvestmen
Pettalidae, confirming previous work (Boyer & Giribet, 2007;
Boyer et al., 2007b). Aoraki, Neopurcellia and Rakaia are not
each other’s closest relatives (Figs 2, 5–7). Therefore, we
conclude that the present-day diversity of Cyphophthalmi
found in New Zealand does not represent a single radiation,
but in fact three separate evolutionary lineages.
The lack of resolution among pettalid genera from different
land masses prevents the use of a molecular clock in assigning
dates to nodes on this particular phylogeny. In the absence of
divergence estimates, what can these results tell us about
dispersal vs. vicariance within the Gondwanan family Pettalidae? In a previous phylogenetic study of Cyphophthalmi, the
age of the family Pettalidae was estimated to be between 178
and 215 Ma, with its origin in the Jurassic/Triassic (Boyer
et al., 2007b). As New Zealand was the last major land mass to
separate from temperate Gondwana, it is unlikely that the New
Zealand genera have originated in the past 25 Ma, while all the
other genera have been in place for more than 100 Ma – and
that none are paraphyletic – as would be the case under a postOligocene drowning dispersal scenario.
If New Zealand has indeed been submerged, and the
terrestrial biota of the archipelago has arrived solely through
transoceanic dispersal, we would expect to see close relationships between the pettalid fauna of New Zealand and species
from other nearby land masses. In many groups of plants, New
Zealand representatives have clearly dispersed from Australia
(Gibbs, 2006; Sanmartı́n, 2007). In some of our analyses, the
monophyletic genus Austropurcellia from Queensland appears
as the sister taxon of Rakaia, but Rakaia never appears as a
lineage nested within a paraphyletic Austropurcellia, as one
would expect under a scenario of dispersal from eastern
Australia. Some elements of the New Zealand biota, such as
parakeets, have been postulated to have arrived from New
Caledonia (Gibbs, 2006). In the case of mite harvestmen,
dispersal to and from New Caledonia is strongly rejected,
because the mite harvestmen of New Caledonia belong to a
different family endemic to that island, Troglosironidae, which
is in turn closely related to the Tropical Gondwanan family
Neogoveidae from tropical West Africa and tropical South
America, and not to Pettalidae (Boyer et al., 2007b). Although
dispersal has certainly played a role in establishing some
elements of the New Zealand biota, and although it is not
possible to unequivocally reject dispersal in the case of mite
harvestmen, our data do not support the evolutionary
relationships expected if we assume dispersal as an explanation
for the presence of the family Pettalidae in the archipelago.
Within New Zealand, the geographical ranges of the three
genera of Cyphophthalmi can be described as follows. The
genus Neopurcellia (sensu Boyer & Giribet, 2007) comprises a
single widespread species, although we do not discard the
possibility of this constituting a series of cryptic species (S.L.B.,
unpublished data) – found throughout the west coast of the
South Island as far north as Lake Kaniere, and as far south as
Te Anau (Fig. 2). The genus Rakaia (sensu Boyer & Giribet,
2007) is found on Stewart Island, along the south and east
coasts of the South Island and throughout Nelson, and in the
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
North Island as far north as Lake Waikaremoana (Fig. 2).
Rakaia does not overlap in distribution with Neopurcellia, but
certain lineages of Rakaia do overlap with Aoraki in the
Nelson/Marlborough area and in the North Island. Finally, the
genus Aoraki is found throughout the North Island as well as
in the South Island as far south as Aoraki/Mt Cook. It is not
found on Stewart Island, or anywhere along the south and east
coasts of the South Island, where Rakaia is highly diverse
(Fig. 2).
Taken together, the distributions of these genera indicate
two major trends. Firstly, the Southern Alps form a barrier not
readily crossed by mite harvestmen. Only Neopurcellia inhabits
the very wet forests to the west of this mountain range, except
in the most northern reaches of the Southern Alps, where
certain members of both Aoraki and Rakaia are found.
Secondly, Aoraki and certain lineages of Rakaia have diversified within an overlapping range, including the North Island
plus the northern part of the South Island. Close relationships
between North and South Island clades have also been
observed in other groups of animals, such as brown kiwis
and cicadas (Baker et al., 1995; Buckley et al., 2001). These two
land masses were connected by a large region of grassland and
shrubland during the height of the Pleistocene glaciation only
20,000 years ago (McGlone et al., 1993).
The South Island: west coast, Southland, Otago and
Canterbury
The two most widespread species of Cyphophthalmi in New
Zealand are Aoraki denticulata (Boyer et al., 2007a) and
N. salmoni. Each of these species occurs in the extensive
Nothofagus and podocarp forests of the western South Island,
where rainfall is extremely high. These are the largest ranges of
continuous forest in the South Island, and such large ranges of
these two species no doubt reflect the extensive suitable habitat
in which they live.
The Southern Alps mountain range has created a barrier that
divides the high-rainfall western part of the South Island from
the dry eastern part. Forests to the east of the Southern Alps
are much patchier and drier than those to the west, possibly
driving allopatric speciation in this area. There are three closely
related species in the Rakaia clade b with adjacent but nonoverlapping distributional ranges in the south coast of the
South Island, and four closely related species from Rakaia
clade c with adjacent but (again) non-overlapping ranges in
the central eastern part of the South Island (Fig. 5). In contrast
to the large, continuous forests in the north and west parts of
the South Island, which are home to widespread species, the
patchy, discontinuous forests along the east coast of the South
Island are home to species with very restricted ranges.
Nelson, Marlborough and the North Island
There is a close relationship between Cyphophthalmi from
Nelson/Marlborough and the North Island in both Aoraki and
Rakaia – with several localities home to pairs of species from
1095
S. L. Boyer and G. Giribet
the two genera. A. granulosa and R. media, A. inerma and
R. media, A. denticulata and R. magna australis, A. denticulata
and R. florensis, and A. denticulata and R. minutissima are all
pairs of taxa which we have found living sympatrically at single
collecting sites. Although the Cook Strait today constitutes
a formidable barrier for mite harvestmen to cross, only
20,000 years ago, at the height of the glacial maximum, the
Nelson/Marlborough area was connected to the North Island
by an area of grassland and shrubland. During this time,
scattered forests were present throughout the North Island as
well as in the northernmost parts of Nelson and Marlborough.
It is therefore plausible that Cyphophthalmi persisted across
this area during the Pleistocene.
Stewart Island
Within Rakaia, the two species from Stewart Island and two
species from Nelson + North Island form a clade (Rakaia clade
a), which is sister to all other species in the genus. This
relationship among the Rakaia clade a species is one of the
most stable and well supported clades within the New Zealand
Cyphophthalmi; it is found in the strict consensus of all the
shortest trees under all parameter sets for the combined
molecular data set, as well as in the strict consensus of all
shortest trees under all parameter sets for the 18S rRNA, 28S
rRNA and histone H3 data partitions. Bayesian analyses of
every data partition retrieved this clade with > 95% posterior
probability, and this relationship was also found in the optimal
trees of every likelihood analysis and in our previous study of
the family Pettalidae incorporating molecular data (Boyer &
Giribet, 2007).
From a morphological point of view, this relationship is not
surprising. Two of these species (R. florensis and R. minutissima) have a fully bisegmented male tarsus IV, a condition
which is probably plesiomorphic in Pettalidae, as Parapurcellia
and Neopurcellia share this character. The two species sister to
R. florensis and R. minutissima have male tarsi IV, which
display partial bisegmentation apparent in both light microscope and SEM images (see fig. 7d–i in Boyer & Giribet, 2007).
Rakaia minutissima and R. stewartiensis also share an anal plate
with a highly concave posterior margin not seen in any other
New Zealand Cyphophthalmi (see figs 8 & 9 in Boyer &
Giribet, 2007).
Although a close relationship between animals from
the southernmost South Island and the northern South
Island + southern North Island may seem counterintuitive,
similar patterns of disjunction are common across many
groups of New Zealand plants and animals (Heads, 1998;
Trewick & Wallis, 2001; Heads & Craw, 2004; Haase et al.,
2007), and two alternative hypotheses have been proposed to
explain this pattern. One possibility is that this disjunction
reflects the relicts of groups that were once more widespread.
During the LGM, ice covered much of the south-west coast of
the South Island, comprising the area that would have linked
Stewart Island to the regions currently inhabited by R. florensis
and R. minutissima. This could have extirpated Cyp1096
hophthalmi inhabiting the area, leaving intact only the Stewart
Island and Nelson + North Island members of what was once
a much more widespread clade. Another possible explanation
for South Island disjunctions is displacement along the Alpine
Fault, a hypothesis that is supported in a study of hydrobiid
gastropods (Haase et al., 2007) but not across larger and more
mobile terrestrial arthropods (Trewick, 2001).
CONCLUSIONS
The mite harvestmen (Cyphophthalmi) fauna of New Zealand
includes three evolutionary lineages, which clearly are members of the temperate Gondwanan family Pettalidae, and which
are not closely related to each other. The relationships of the
New Zealand genera to relatives in Australia and other adjacent
land masses, such as New Caledonia, provide no evidence for
recent dispersal as the explanation for the presence of pettalids
in New Zealand. In the southern regions of the South Island,
Neopurcellia occurs to the west of the Southern Alps, while
Rakaia occurs to the east of these mountains. In the North
Island, the genus Aoraki predominates. The latter two genera
overlap in the southern part of the North Island and in the
northern region of the South Island, reflecting the recent
historical connection of these two major islands. Despite the
lack of precise dating, the history of Cyphophthalmi in New
Zealand has clearly been influenced by ancient Gondwanan
vicariance, as well as more recent alterations to the New
Zealand landscape during the Pliocene uplift of the Southern
Alps and Pleistocene glaciation across the South Island. Future
comparative studies of phylogeographical relationships within
species from disparate geographical regions of New Zealand,
such as the South Island’s west coast and east coast, will
increase our understanding of the role that habitat fragmentation has played in the diversification of this remarkable
group of short-range endemics.
ACKNOWLEDGEMENTS
For help with field work, the authors acknowledge Greg
Edgecombe, Cyrille D’Haese, Jessica Baker, Phil Sirvid, Ricardo
Palma and Cor Vink. All permits were arranged with the help
of the sublime New Zealand Department of Conservation;
special thanks go to Ian Millar at the Nelson office. Phil Sirvid
and Ricardo Palma at Te Papa Tongarewa/Museum of New
Zealand (Wellington) loaned holotype materials as well as
other specimens that provided valuable data regarding distributions of species; some of these materials included the
Department of Conservation’s pitfall trap specimens for Aoraki
inerma, A. tumidata and Rakaia antipodiana. Additional
specimens were loaned by Simon Pollard from the Canterbury
Museum (Christchurch), Abigail Blair from the Otago
Museum (Dunedin), and Grace Hall at the New Zealand
Arthropod Collection of Manaaki Whenua Landcare Research
(Auckland). The authors would like to thank Maureen O’Leary
for the invitation to participate in the MorpoBank project;
Jennifer Lee assisted with the uploading of images to the
Journal of Biogeography 36, 1084–1099
ª 2009 Blackwell Publishing Ltd
Biogeography of New Zealand’s mite harvestmen
database. Emily Sabo assisted with the production of Fig. 2.
This material is based on work supported by the National
Science Foundation under Grants Nos 0236871 and DEB0508789. Field work for this project was also generously
supported by a Museum of Comparative Zoology Putnam
Expedition Grant.
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SUPPORTING INFORMATION
Additional Supporting Information may be found in the
online version of this article:
Appendix S1 Collection information, voucher numbers, and
GenBank accession numbers for the specimens used in this
study.
Appendix S2 Morphological matrix.
Please note: Wiley-Blackwell is not responsible for the
content or functionality of any supporting materials supplied
by the authors. Any queries (other than missing material)
should be directed to the corresponding author for the article.
BIOSKETCHES
Sarah Boyer earned her PhD in Biology from Harvard
University and is Assistant Professor of Biology at Macalester
College. She is interested in the phylogeny, biogeography and
phylogeography of terrestrial and freshwater invertebrates. A
particular focus of her research is the fauna of New Zealand.
Gonzalo Giribet is Professor of Organismic and Evolutionary Biology and Curator of Invertebrates at the Museum of
Comparative Zoology, Harvard University. He is interested in
the origins and maintenance of invertebrate diversity, in both
marine and terrestrial environments, and in theoretical aspects
of systematics and biogeography.
Editor: Malte Ebach
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