awc newsletter sept, 2006
TRANSCRIPT
1
Figure 1: Māori variety of bottle gourd (hue) from the Auckland region, growing in
Otaki, Kapiti Coast during late summer, 1999. (Photo credit: Mike Burtenshaw).
Issue Number 4 (September 2006)
Origins and Dispersal of the Polynesian Bottle Gourd
Our research group at the Allan Wilson
Centre have discovered the bottle
gourd (or hue in Māori) grown in
Polynesia originated in both Asia and
the Americas. The bottle gourd, which
is closely related to the pumpkin, is one
of the many crops that Polynesians
took with them as they settled the
islands of the Pacific, including
Aotearoa New Zealand.
Anthropologists had previously
suggested the bottle gourd had come
from South America along with the
sweet potato (kumara), but our
research shows there is also a
significant genetic contribution from
Asia, and that Polynesian bottle gourds
are in fact hybrids between gourds from
both of these continents.
We collected a large number of bottle
gourds seeds from Asia and the
Americas, as well as eight Māori bottle
gourds from New Zealand.
Ngā Orokohanga me ngā Tuaritanga o te Hue o Te Moana nui a Kiwa
Kua kitea e tō mātou rōpū i te Allan
Wilson Centre, ko te Hue (he Bottle
Gourd i te reo Ingarihi) e tipu ana ki Te
Moana nui a Kiwa, i taketake mai i Āhia
me ngā motu o Amerika. Ko te hue
tētahi o ngā huawhenua maha i kawea
mai e ngā tāngata o Te Moana nui ā
Kiwa i a rātou e noho haere ana i ngā
moutere o te Moana nui ā Kiwa, tae
noa ki Aotearoa. He whanaunga tata te
hue ki te paukena.
I ngā rā ki muri, i kī ngā tohunga
tikanga tangata i taketake kē mai te
hue i Amerika ki te Tonga i te taha o te
kūmara, heoi kua kitea i roto i tō mātou
rangahau, te kaha uru o ngā momo
whakaheke mai i Āhia ā, ko te mea kē,
he kākano whakauru kē nō ngā motu e
rua nei.
He tino maha ngā kākano hue i
kohikohia e mātou i Āhia me ngā wāhi
o Amerika whānui, tae noa ki ngā hue e
waru o Aotearoa.
Inside this issue
Origins and Dispersal of the Polynesian Bottle Gourd ..........1 Ngā Orokohanga me ngā Tuaritanga o te Hue o Te Moana nui a Kiwa ...................................1
‘Giant’ Collembola of New Zealand: The Largest Springtails in the World!................................4
Tuatara Assisting with Education Outreach .....................................7 Celebration of Te Kopinga, First Marae of the Moriori ..................8 Phylogeography of Carnivorous Land Snails (Family Rhytididae) ................10 Recent Publications ................14 Contact Us ................................16
TThhee AAllllaann WWiillssoonn CCeennttrree NNeewwsslleetttteerr
The Māori gourds were obtained from marae and heritage
seed companies, and are thought to be derived from true
Māori bottle gourds grown in pre-European New Zealand.
This collection was used to develop DNA markers that
could be used to trace the gourd’s origins. We used DNA
fingerprinting, similar to that used to identify humans, to
locate regions of the gourd genome that are variable. Just
as in humans, individual bottle gourds share nearly
identical DNA – probably more than 99% – so the DNA
fingerprinting is used to identify the less than 1% of the
DNA that makes each bottle gourd different. These
variable DNA fragments could then be used as DNA
markers to trace the origins of the Polynesian bottle gourd.
The DNA markers showed that Asian gourds are all of one
type, American gourds are all of another type, and that
Polynesian gourds are a mixture of both. This opens a
number of possibilities for the dispersal of this species.
I tīkina mai ngā hue Māori i ngā marae me ētahi kamupene
pupuri ā-tikanga i ngā kākano ā, ko te whakaaro, i ahu mai
ēnei i ngā hue a te Māori i whakatipuria i mua i te taenga mai
o te Pākehā.
I whakamahia tēnei kohikohinga hei hanga tohu pītau-ira
(DNA) hei whakataki i te takenga mai o te hue. I whakamahia
e mātou te tapukara pītau ira (DNA) rite ana ki tērā e
whakamahia ana ki te tautuhi i te tangata, hei rapu i ngā wāhi
tipu ai te hue whai tāupe. He tino ōrite katoa nei ngā pītau-ira
o ia hue, pērā anō ki te tangata – te āhua nei nui atu i te 99
ōrau – nā reira ka whakamahia te tapukara pītau-ira hei
tautuhi i te toenga o te 1 ōrau o te pītau-ira, e rerekē ai tēnā
hue ki tēnā hue. Ka taea ēnei maramara pītau-ira tāupe te
whakamahi hei kai tohu pītau-ira, hei whakataki hoki i te
takenga mai o te hue o Te Moana nui a Kiwa.
I whakaaturia mai e ngā kaitohutohu pītau-ira he momo kotahi
ngā hue katoa o Āhia, he momo kotahi atu anō ngā hue o
Amerika, ā he raranu o ngā mea e rua te hue o Te Moana nui
ā Kiwa. Nā konei, kua puta ngā whakaaro mō te puananī o
tēnei momo.
Figure 2: Pai Kanohi with gourd containers (tahā huahua) for preserving wood pigeons (kererū), circa
1910. Ruatahuna, Huiarau Range (just north of Lake Waikaremoana), North Island.
(Photo credit: Archives New Zealand and the Alexander Turnbull Library, Wellington).
2
Bottle gourds could have been brought from Asia with the
ancestors of Polynesians when they moved out of South
East Asia 5,000 years ago, or perhaps with later migrants
from Asia. The American gourds could have been
introduced to Polynesia with the kumara. Polynesian
voyagers are thought to have sailed from Easter Island to
South America about 1,000 years ago and collected the
kumara before sailing back to Polynesia. The bottle gourd
is also very buoyant, so we cannot rule out that a gourd
floated from Asia or the Americas to Polynesia, where it
was picked up from a beach and propagated from the
seeds which are stored inside the fruit.
The bottle gourd was one of the most important crop
species in pre-European Polynesia. In New Zealand young
bottle gourds were eaten (like zucchini), but were mainly
used when dry and mature. These hard-shelled bottle
gourds were hollowed out and used primarily as water-
carrying vessels, containers for food (muttonbirds and tui
were stored in their own fat), musical instruments and
canoe bailers. Māori bottle gourds are still grown in New
Zealand today, but mostly for ornamental purposes (such
as the one pictured) and to preserve this important part of
Polynesian and Māori life.
Acknowledgement: We are grateful to Mr Jonathan
Procter and Rangitaane O
Manawatu for supporting the
genetic work undertaken on the
bottle gourd.
Tērā pea i haria mai te hue i Āhia e ngā tūpuna o Te Moana
nui ā Kiwa i te wā i puta ai rātou i Āhia ki te tonga, e 5,000 tau
ki muri. Tērā pea i haria mai ngā hue Amerikana ki Te Moana
nui ā Kiwa i te taha o te kūmara: Inā hoki, tērā tētahi kōrero, i
haere ngā kaiwhakatere waka o Te Moana nui ā Kiwa mai i
Rapanui ki Amerika ki te Tonga, he āhua 1,000 tau ki muri, i
reira kohikohi ai i te kumara i mua i tōna hokinga ki te Moana-
nui-Kiwa. He tino māngi te hue, nā reira ko wai ka mōhio tērā
pea i māunu kē mai i Āhia, mai i ngā wāhi o Amerika rānei, ki
Te Moana nui ā Kiwa. I reira ka kohia mai te ākau ka
whakamakuru ai ngā kākano o roto i te hue.
Ko te hue tētahi o ngā momo huawhenua tino hira i mua i te
taenga o ngāi Pākehā ki Te Moana nui ā Kiwa. I kainga ngā
hue iti (pērā ki te zucchini) engari i tino whakamahia ngā hue i
te wā kua hua, kua maroke hoki. I hākarohia ēnei hue mārō
nei ana, kātahi ka whakamahia hei oko kawe wai, hei ipu, hei
kūmete kai rānei (ka huahuatia ngā tītī me ngā tūi) hei taonga
pūoro, hei tīheru mō te waka. E whakatipua tonu ana te hue
a te Māori, i Aotearoa nei, heoi hei whakapaipai noa iho,
(pērā ki tā te pikitia nei) i te nuinga o te wā, me te whakaora
tonu i tēnei āhuatanga o Te Moana nui ā Kiwa me te ao o te
Māori.
NGĀ MIHI
Ngā mihi ki a Jonathan Procter me Rangitāne
o Manawatū mō tā rātou tautoko i ngā mahi ira
tangata i whakahaerehia e pā ana ki te hue.
Figure 3: Ornamental bottle gourd carved with
modern Māori design. (Photo credit: Andrew Clarke).
Andrew Clarke PhD student Massey University [email protected]
English to Māori translation by Māori Language Services, Māori Language Commission – Te Taura Whiri i te Reo Māori.
3
‘Giant’ Collembola of New Zealand: The Largest Springtails in the World!
What do Collembola do?
4
Collembola (springtails) are an ancient
(>412MYA) and highly successful class
of hexapod dating back to at least the
Devonian (Rhyniella praecursor) or
Upper Silurian. Although predominantly
soil and litter dwellers, they also occur
in a wide range of habitats such as on
vegetation, under rocks, in logs (Fig. 1),
in tree canopies, in caves, in the marine
littoral zone, and in freshwater systems.
Figure 1: A typical habitat within a South Island (Arthurs
Pass) beech forest. (Photo credit: Mark Stevens).
Figure 2: Collembola (Holacanthella duospinosa.) collected
from Ohakune. Known to reach 17mm in length, which
makes it the largest known springtail in the world.
(Photo credit: Rod Morris).
As detritivores, springtails are an
important group in nutrient cycling and
are beneficial organisms as very few
species feed on live plant material. The
ecology and widespread nature of
springtails suggest that they warrant
more attention from biologists.
Worldwide, over 7000 species in 581
genera have been described and are
found throughout the world including
the Arctic and Antarctic regions.
The ‘Giants’
The most spectacular and largest
springtails form the subfamily
Uchidanurinae Salmon, 1964. The
Uchidanurinae currently consists of
eight genera and 15 species all of
which are endemic to their respective
localities—China (Assamanura
besucheti), Indo-China (Denisimeria
caudata, D. longilobata, D. martyni),
Micronesia/Polynesia (Uchidanura
bellingeri, U. esakii), New Caledonia
(Caledonimeria mirabilis), eastern
Australia/Tasmania (Megalanura
tasmaniae, Acanthanura dendyi,
Womersleymeria bicornis), and New
Zealand (Holacanthella spinosa, H.
paucispinosa, H. brevispinosa, H.
duospinosa, H. laterospinosa).
These species are particularly
remarkable in that some are the largest
springtails recorded world-wide (up to
17 mm long for the New Zealand
species), and most sport coloured
digitations (spine-like projections) on
their dorsal and lateral surfaces (Fig.
2), and are saproxylic (live within
decomposing logs).
Saproxylic Communities
Saproxylic communities drive
nutrient cycling and nutrient
uptake by plants in forests. This
action returns nutrients locked up
in dead wood to the ecosystem
where they support large and
diverse invertebrate populations
and enrich the soil to enhance
growth and regeneration. A large
proportion of the New Zealand
endemic plants and animals
considered to be of conservation
importance are adapted to native
forests and saproxylic
communities are an important part
of these ecosystems. As well as
enriching forest soils, saproxylic
organisms (which include, for
example, earthworms, myriapods,
fungi, beetles and spiders) provide
important food sources (directly
and indirectly) for a number of
New Zealand’s most treasured
and threatened species including
Kiwi, rhytidid snails (including the
Powelliphanta), robins and velvet
worms (Peripatus), but the
Uchidanurinae are currently only
considered to be of extreme
conservation status in Australia. They
are likely to be a particularly important
part of New Zealand’s saproxylic fauna
as springtails have been shown to be
key agents in controlling the dynamics
of soil microorganisms (bacteria, fungi
and algae), and thus play a crucial role
in defining the composition of the
saproxylic community.
What are we doing?
Despite the overwhelming ecological
importance of New Zealand’s
Uchidanurinae they have not been the
subject of scientific interest since their
original descriptions between 1899 and
1944. In recent times the forests they
inhabit have undergone large scale
fragmentation following first Polynesian,
then European settlement, and
subsequent infestation by introduced
pests. Future scientific and/or
conservation effort requires a greater
understanding of their distribution, but
with only a total of eleven historical
records determining what effect
disturbances have had on these unique
and important springtails has been an
arduous task.
Our work aims to:
(1) Provide a detailed examination of
the distribution of all five Holacanthella
species throughout New Zealand.
(2) Develop an updated key to their
identification (available online:
http://awcmee.massey.ac.nz/people/ms
tevens/NZ.htm)
(3) Examine phylogenetic relationships
for the New Zealand, Australian and
New Caledonian species using
mitochondrial and nuclear DNA.
(4) Examine the phylogeographic
patterns for the three widespread New
Zealand species (H. brevispinosa, H.
paucispinosa, H. duospinosa) using
mitochondrial and anonymous nuclear
markers.
Distribution of all five Holacanthella species throughout New Zealand
Throughout New Zealand the density of
Holacanthella individuals found at any
particular site was highest in beech
forests (Nothofagus spp.), and lower in
Figure 3: Sampling for Collembola in rotting logs in Hawdon Valley, Arthurs Pass. Robins are frequent visitors (bottom right)
making the most of a free feed! H. paucispinosa (top left), H. spinosa, (middle) and H. duospinosa (bottom) are among the many
Collembola found here. There is still very little known about the organisms which make up part of the saproxylic community. (Photo credits: Mark Stevens and Rod Morris).
5
6
other mixed forest types. Several sites
possessed more than one species, for
example locations in southland,
Fiordland, Arthurs Pass, Lewis Pass,
Mt Arthur Tableland, Wellington, and
Ohakune. The distribution of the two
species H. brevispinosa and H.
paucispinosa was almost completely
overlapping (sympatric) extending
from Stewart Island, throughout South
Island, and extended north to the
Central Plateau of North Island.
Finding both species in or under a
single log occurred on several
occasions. Holacanthella spinosa is
the only other South Island species. At
Mt Ruapehu, on the Central Plateau,
the three species were found together
with another species, H. duospinosa,
and this species extends north to
Northland (including Kawau I., Little
Barrier I., Great Barrier I.). The fifth
remaining species, H. laterospinosa, is
only known from Cuvier Island off the
Coromandel Peninsula (North Island).
Figure 4: Holacanthella paucispinosa collected from Rahu
Saddle, South Island. (Photo credit: Rod Morris).
With a recent summer student (David
Winter) and numerous field helpers we
have extended the known distribution of
all five New Zealand endemic
Holacathella species. The historic
(MONZ) and new records highlight the
importance of maintaining old growth
forests in the west coast and northern
South Island, central North Island, and
Cuvier Island to adequately preserve
these species. Molecular and
morphological studies are now
underway to further examine the
intraspecific (within species)
morphological variability that we
observed across the ranges for these
species.
Conserving forgotten species
The loss of habitat emphasises human
impacts which is currently likely to be
the greatest threat to this group. Most
importantly, available dead wood on the
forest floor is a requirement of these
saproxylic communities. ‘Natural’
forests (unmanaged) currently support
large populations of Holacathella. Most
notable are beech forests that have not
undergone extensive logging, such as
in southland, Abel Tasman National
Park (Mt Arthur tableland), and the
Tongariro National Park (Central
Plateau), all support dense, species
rich populations. However, most of New
Zealand’s remnant forests are broken
into small fragments. In total there are
around 120,000 such fragments with an
average size of 53.9 Ha. Collembola
are known to be highly sensitive to
forest practices and their low dispersal
capacity makes recolonisation of
disturbed (and regenerating) sites more
difficult, particularly if these are
fragmented. The preservation of the
‘natural’ characteristics of these
habitats and their original species
composition appears essential.
Holacanthella are an under-studied
group that are likely to be an important
part of New Zealand’s forgotten
invertebrate biota. At present the
Department of Conservation
Invertebrate recovery plan makes no
mention of any of New Zealand’s
springtail species. Collembola are
typically considered too small and too
numerous to be considered in need of
conservation. However, this is not
always the case: a reserve in
Tasmania (Springtail Reserve) has
been dedicated solely for a species of
Collembola, Tasphorura vesiculata;
species of Uchidanurinae were listed by
the IUCN in the Red Data book in 1994;
and the removal of dead wood is listed
as a threatening process in NSW,
Australia. The likely ecological
importance and the vulnerability of
Holacanthella means they should form
a part of future conservation plans.
Understanding of their distribution and
genetic diversity will aid in determining
vulnerable/rare species and regions.
Our objective is to understand more
fully ‘the small things that run the world’
and the processes that have shaped
New Zealand’s biodiversity.
For further reading:
Collins Field Guide to New Zealand Wildlife.
By Terence Lindsey and Rod Morris. Page
187. ISBN 1-86950-300-7
Mark StevensPostdoc, Massey University, Palmerston North [email protected]
7
Project on Tuatara conservation throughout schools in
New Zealand. (Photo credits: Sue Keall).
Tuatara Assisting with Education Outreach
“Tuatara: a Taonga for the People of
New Zealand”, a joint project between
Victoria University of Wellington, Te
Atiawa iwi, The Allan Wilson Centre, and
the San Diego Zoo was successfully
completed in December 2005. Funded
by the Royal Society of New Zealand’s
Science and Technology Promotion
Fund, this project took conservation
education outreach about tuatara to
schools around New Zealand. An
additional goal of the project was to
provide training to iwi members in tuatara
biology, research and conservation
education.
Training began when two iwi
representatives attended a Conservation
Education Workshop hosted by the San
Diego Zoo in October 2004. The next
step was for them to participate in a
Victoria University research field trip to
North Brother Island in March 2005.
Here they gained first-hand knowledge of
tuatara biology and behaviour, and
learned techniques in scientific research.
In April there was a week long visit to
Victoria University in which the two
teachers built on their knowledge of
tuatara biology, the results of scientific
research and how it is being applied to
tuatara conservation. Several
Wellington primary schools were visited
so that trial presentations could be
given. A 30 minute narrated
Powerpoint presentation conveyed how
science and technology play an
essential role in supporting the
conservation of native biodiversity. The
presentation concluded with a live
tuatara being shown to attendees on an
individual basis, and in most cases
touching the tuatara was encouraged.
Once training was complete, the project
visited schools in five centres around
New Zealand during 2005 –
Blenheim/Picton, New Plymouth,
Whakatane, Whangarei and
Greymouth. Presentations were given
at 57 primary schools, 12 secondary
schools and 9 public venues. Each
school group consisted of 50 students
and several teachers (limited in size for
the welfare of the tuatara): public
groups ranged in size up to 100.
Approximately 3500 members of the
New Zealand public participated in the
programme in total. The presentations
were enthusiastically received, and the
opportunity to meet and touch a live
tuatara had real impact. Substantial
feedback about the educational value
of the presentation was received, with
extremely positive comments such as
“a fabulous presentation and one which
the students will remember always”.
Media interest was also high, with 19
newspaper articles reporting the
project’s school and public
presentations.
The project has enabled iwi presenters
to develop knowledge and skills that
will assist them in developing further
conservation education outreach
programmes within their own rohe. An
additional outcome has been the
positive example set by these young iwi
teachers to their peers, as to what can
be achieved with further training in
science and conservation of our taonga.
Sue Keall Technical Officer Victoria University of Wellington
Celebration of Te Kopinga, First Marae of the Moriori
Last year I was given a very special
opportunity, to attend the opening
celebration of the first Moriori marae on
Rekohu/ Chatham Island. I was invited
because of my involvement with the
Moriori (indigenous people) through my
research.
8
My PhD is on the conservation genetics
of the Chatham Island Taiko, called
Tchaik by Moriori. This petrel is New
Zealand’s most endangered seabird. In
the past it was an important food
source for some Moriori imi (iwi/tribes).
The birding practice was highly
ritualised and involved special karakii
(karakia/prayers). Conservation was
important to the Moriori and they were
very careful not to destroy burrows
when collecting chicks. However, once
predators were introduced, the Taiko
population was quickly decimated and
no longer a viable food source. The last
recorded birding trip happened in 1903
when 300 chicks were taken. The Taiko
remains a taonga (treasured) species
to the indigenous people.
Figure 1. View of Te Awapatiki, mouth of Te Whanga (lagoon), ancient meeting place of the Moriori. (Photo credit: Hayley Lawrence).
The Moriori population was also
devastated around the same time as
the Taiko population, when other
peoples invaded the Chatham Islands.
Outside of the Chathams, the Taiko
was thought to be extinct as were the
Moriori people themselves, but the
Moriori and the Taiko did survive. The
Taiko was rediscovered on the night of
New Year’s day 1978, by David
Crockett. Mainstream New Zealand
became aware of Moriori survival when
a documentary was filmed in 1980.
After this, the Moriori people
commissioned a book by Michael King
about their history, language and
culture. A Waitangi Tribunal claim in
2001 brought official recognition of their
unique status. The opening of the first
Moriori marae was a great achievement
in the reaffirmation of culture and the
joining together of Moriori people.
Te Kopinga is the first Moriori marae
because instead of marae, Moriori used
to meet in groves of Kopi (Karaka)
trees. The new marae looks over Te
Awapatiki, the mouth of the lagoon,
where all imi met in the past. The
building complex of the wharenui (main
house), kitchen, dining hall, and
administration blocks, was designed so
that by air it looks like an albatross in
flight. The albatross is also a taonga
species to the Moriori. “Hokomenetai” is
the name of the wharenui, a house of
peace. It is in a pentagonal shape
emulating the rocks in the basalt
columns found on the island.
Around 1000 people were present at
the opening celebrations. A dawn
ceremony began the day, very
appropriate since the Chathams is the
first place to see the sun. At lunchtime
the official whakamaurahiri (welcoming)
began. It was slightly different from a
Maori powhiri. Moriori Kuia called the
maurahiri (manuhiri/visitors) on to the
marae while the Ratana church band
led us. We then entered the wharenui
for the hau-rongo (speeches). There
was no wero (challenge) because the
Moriori live under Nunuku’s law, a
covenant of peace. Karakii were said
as Ka Pou o Rangitokona (the central
post) was blessed. Moriori Rangata
Matua (Kaumatua/elders) and leaders
spoke, some in Moriori, some in Maori,
and some in English. Moriori rongo
(waiata/songs) were sung after each
speaker. Speeches were made by
invited guests, including Kaumatua
from Maori iwi from across Aotearoa
and Te Wai Pounamu (NZ). Michael
King’s son spoke in his honour,
followed by Helen Clarke. At the
conclusion of the speeches, Moriori
children, and Maori and Pakeha
children from the island renewed the
covenant of peace. During this moving
ceremony, Moana and the tribe (the
band) sang a song specially composed
for the occasion.
After the ceremony was kai time. The
feast was amazing and included a
bounty of kaimoana (seafood). People
exclaimed in delight at the size of the
koura (crayfish) that the Chathams is
renowned for. Other delicacies included
akoako (titi/muttonbird), hangi, and
weka (a bird which can only legally be
eaten on the island). After the feed,
Moana and the tribe (formerly Moana
and the Moahunters) entertained us.
After that many people headed off to
the legendary Hotel Chathams, which
has its own Island Gold beer, White
Pointer vodka, and Blind Jims bourbon.
(I was warned about the bourbon!)
The trip was fun, but also useful for my
project. Imi/Iwi consultation is essential
for a few aspects of my work, including
cloning and bone collecting. I believe it
is important to establish good
relationships with imi/iwi, especially
when the species you are working on is
a taonga to them. The relationship can
be very rewarding for both sides.
During my trip, I reaffirmed old contacts
9
Figure 2. Hokomenetai, the wharenui of Te Kopinga marae. (Photo credits for this page: Hayley Lawrence).
and made new ones. The organisers
genuinely thanked me for coming,
which surprised me because I felt
honoured just to be asked. I think that
they appreciated my attendance as
demonstrating that our relationship is
important to me. I distributed
information to interested people and
displayed a poster. It included a
request for information regarding
traditional ecological knowledge, but
also included an invitation for people to
contact me if they would like to know
more about my work (reciprocity is
important). (The poster is also available
on the AWC website at:
http://awcmee.massey.ac.nz/project_H
Lawrence.htm)
I also visited other people I know in the
Taiko Trust and Department of
Conservation. They have helped me
greatly with my project, especially
logistically, for which I am very thankful.
Getting to know them on a personal
level has been great. All in all, my visit
to Rekohu was a very rewarding
experience. I am grateful to IMBS and
AWC for realising the importance of this
trip and providing funding for it, and to
the Hokotehi Moriori Trust for inviting
me.
Nau te rourou,
Naku te rourou,
Ka ora ai te iwi … a, mo tenei kaupapa,
ka ora ai te Taiko
Hayley LawrencePhD student Massey University - Albany [email protected]
Phylogeography of Carnivorous Land Snails (Family Rhytididae)
10
Figure 2: Amborhytida dunniae. (Photo credit: Fred Brook).
New Zealand Rhytididae
The New Zealand Rhytididae are a
large group of carnivorous land snails
that include the well known
Powelliphanta group. Members of this
large family can also be physically very
large – at up to 90 mm some
Powelliphanta are New Zealand’s
largest land snails. Other members of
the group, like Paryphanta, may get up
to 75 mm. With spectacular shells,
these snails have a Gondwanan
distribution: that is, they are found
throughout New Zealand, Australia,
New Guinea, and South Africa.
Figure 1: Payphanta busbyi.
(Photo credit: A. M. Spurgeon, supplied from
New Zealand Mollusca website:
http://www.mollusca.co.nz/)
Rhytidids are carnivorous on worms, as
well as other snails and slugs.
Although these are our largest and
perhaps most charismatic land snails,
their classification is still relatively
poorly understood, and many
of the group’s members are of
conservation concern (mostly
due to rat predation and
habitat destruction).
To be able to address issues
about the conservation status
of the members of this group,
we first need to be sure what
taxonomic groups we are
actually dealing with. One
way in which we can do this is
to compare the results from using
molecular markers with the
expectations you would have from
morphology. By utilizing sequence
data we can evaluate any potential
classification problems there may be
due to either conserved morphologies
or rapidly changing morphological
characters (or a combination of these).
While investigating the classification of
these snails is a useful and worthwhile
purpose on its own, these studies can
be put into context by investigating the
evolutionary history of the groups –
including looking at their
phylogeography.
The Paryphantinae: the Kauri Snails and Relatives
The first study of New Zealand
Rhytididae that we have completed is
one on the Kauri snails and their
relatives. In this group most of the
species are restricted to Northland.
Within the Paryphantinae there are four
genera of large species: Paryphanta
(Kauri Snails, found throughout
Northland), Rhytidarex (from the Three
Kings Islands), Amborhytida (found
throughout Northland), and
Schizoglossa (Paua Slugs, found in the
northern half of the North Island).
In this initial study we set out to
investigate the relationships of the taxa
restricted to Northland and to place
those relationships within a geographic
context. Thus we focused on
Paryphanta and Amborhytida, the
genera restricted to, but widespread
within, Northland. The Kauri Snails
contain two species, Paryphanta busbyi
(Figure 1) (found from Awanui to
Warkworth, which grows up to 75 mm)
and Paryphanta watti (found in the Far
North only, growing up to 60 mm).
Amborhytida contains five nominal
species, three of which have
reasonably wide distributions:
Amborhytida dunniae (Figure 2) (found
from Awanui to Auckland), Amborhytida
forsythi (found from Karikari to north
Kaipara, and historically considered
closest to Amborhytida dunniae), and
Amborhytida duplicata (from in the Far
North only).
Questions
Our initial questions included asking,
what are the evolutionary relationships
among these species? What are the
evolutionary relationships among
populations within these species? And
to place it all into some wider context,
do these relationships make sense
geographically and geologically?
Answers
Figure 3: Bayesian tree for the Paryphantinae and outgroups (the support values not shown appear on the magnified versions of the figures [4, 5 and 6]).
To answer these questions we needed
to generate a phylogeny for the group.
We (meaning our collaborator Fred
Brook) collected samples of all the
species of the four paryphantine genera
found in Northland. The samples were
collected from between three and
eighteen locations in Northland. After
the samples reached Otago, we
sequenced an ~1 kb fragment of the
mitochondrial COI gene for each of
them. Various methods were used to
estimate the phylogenetic relationships
of the group – all of which gave very
similar results. The Bayesian tree
produced is shown in Figure 3. This
tree includes a few outgroup taxa (from
the Rhytididae), and shows that all the
paryphantine genera form natural
groups, with Rhytidarex the most basal.
Of the other taxa in this study, the
coverage of the Paua Slugs
(Schizoglossa) was sparse, but they
were included in this tree for
completeness – this genus is currently
the subject of separate, more in depth,
study.
Figure 3 shows that of the main groups
of interest we have a Paryphanta
group, an Amborhytida dunniae group,
and an Amborhytida forsythi group.
The Paryphanta group (Figure 4)
includes P. busbyi and P. watti. The
Amborhytida dunniae group (Figure 5)
includes the nominal species from the
Hen and Chickens, A. tarangaensis,
and Poor Knights, A. pycrofti, and the
morphological variant from Cape Brett,
A. sp. “Motukokako”. The Amborhytida
forsythi group (Figure 6) includes A.
forsythi, A. duplicata and a set of taxa
that were previously called A. forsythi,
but which we are currently referring to
as A. sp. “Aupouri”.
The magnified version of the Bayesian
tree for the Paryphanta group (Figure
4) shows several interesting results.
11
12
Firstly, the phylogeny of Paryphanta
does not correspond with the current
taxonomy of the genus. The two
populations of the Far-North endemic,
P. watti (represented on Figure 4 by
orange stars), that we sampled fell
within a clade including several
Figure 4. Bayesian tree and map of sample locations for the Paryphanta group.
Figure 5. Bayesian tree and map of sample locations for the Amborhytida dunniae group.
populations of P. busbyi (the blue stars
on Figure 4) which extend along the
east of Northland, from near Kaitaia
south to Hen Island and the Waipu
Hills. A second clade included several
populations of P. busbyi (the green
stars on Figure 4) from the western and
southern areas of Northland between
Herekino and north Kaipara, with an
outlying population further south near
Warkworth. There are no obvious
consistent shell differences between
these “eastern” and “western” clades,
whereas shells of P. watti are easily
distinguished from those of P. busbyi:
they have ~1 cm (~15%) smaller
diameter as adults, and have different
colouration. Thus there appear to be
two clades within Paryphanta,
somewhat surprisingly (there is no
simple geological explanation for the
distribution) separated into a
northern/eastern group (blue and
orange stars) and a southern/western
group (green stars).
The magnified version of the Bayesian
tree for the Amborhytida dunniae group
(Figure 5) shows a general lack of
structure (which is at odds with the
structure found in the Paryphanta
group). There is such a lack of
structure in this group that there is no
point in using stars to show the
distribution of different clades within the
group. The morphologically divergent
forms restricted to some of the islands
off the eastern coast of Northland – A.
tarangaensis from Taranga (Hen)
Island, A. pycrofti from the Poor Knights
Islands and A. sp. “Motukokako” from
Motukokako (Piercy Island) and nearby
Cape Brett peninsula – fitted clearly
within the genetic variation ascribed to
A. dunniae. This result suggests that
populations of each of these island (or
near island) endemics are very closely
related, possibly as a consequence of
evolutionarily recent founder events.
The remaining populations of
Amborhytida, originally attributed to A.
forsythi and A. duplicata, formed a
group with very strong support and
were considerably divergent from A.
dunniae (see Figure 3). Thus, the view
of A. forsythi as only subspecifically
distinct from A. dunniae is not tenable,
and in fact the two taxa are locally
microsympatric (e.g., at locations 18
and 19; see Figures 5 and 6).
Moreover, the samples originally
identified from shell morphology as A.
forsythi grouped in a most unexpected
way (Figure 6), falling into two well-
supported non-sister clades, although
the non-sisterhood itself was not well
supported. Populations from Mt Camel,
Karikari Peninsula, and hill country
north of Herekino Harbour,
subsequently referred to in our study as
A. sp. “Aupouri” (the green stars on
Figure 6) were weakly grouped with A.
duplicata (the orange stars on Figure
6), which is endemic to the area
between Cape Maria van Diemen and
North Cape at the northern tip of the
Aupouri Peninsula. Populations of
morphologically similar A. forsythi from
elsewhere in Northland between Taipa
(the type locality) and north Kaipara,
formed a separate, well supported
clade (the blue stars on Figure 6).
The almost simultaneous evolution of
A. duplicata, A. forsythi, and A. sp.
“Aupouri”, which we estimate at being
between 1.9 and 6.6mya, accords with
the inferred former existence of islands
in the Cape Reinga-North Cape, Mt
Camel and Karikari areas during
Pliocene time (1.8–5.3mya). Clearly, A.
duplicata evolved in the Far North and
remained there, with eastern and
western populations subsequently
becoming genetically (but not
conchologically) differentiated over the
last 0.9–3.2my. Possibly, A. sp.
“Aupouri” evolved on what is now
Mount Camel or Karikari Peninsula,
which were also separate islands in the
Pliocene, before spreading south to
Herekino. A. forsythi presumably
evolved in mainland Northland.
What does it all mean?
13
For the Amborhytida forsythi group
(Figure 6) we find interesting and
unexpected patterns, with A. duplicata
falling within the group. This result
raises interesting questions about the
morphological characters that have
previously been used to determine the
relationships within these groups. The
phylogeographic patterns within the
Amborhytida forsythi group make sense
– what makes less sense is that that
they in no way resemble the patterns
within either the Amborhytida dunniae
group or the Paryphanta group.
Because these snails are closely
related, have similar life histories and
live in the same areas, it would have
been reasonable to predict that they
might share similar phylogeographic
patterns (assuming they shared
similar evolutionary histories), but this
is certainly not the case. Whereas the
Amborhytida forsythi group’s
phylogeographic patterns are
relatively straightforward to interpret,
there is no structuring within the
Amborhytida dunniae group, and the
structuring within the Paryphanta
group is incongruous with that of the
Amborhytida forsythi group. Whether
these different patterns (or lack
thereof) represent different ancient
refugial patterns or different abilities to
recolonise different areas after the Figure 6. Bayesian tree and map of sample locations for the Amborhytida forsythi group.
reformation of Northland, or a
combination of these and other
processes, we cannot tell at this point.
What these results do tell us is that if
we had looked at just one of these
groups assuming that, because they
were closely related to one another and
had similar life histories and lived in the
same areas, we could generalise from
one group to another we would have
been very wrong. The discordance in
the phylogeographic patterns in the
groups of snails examined here means
that it is difficult to make strong
inferences about common geological
influences on the evolutionary history of
paryphantines in Northland. If our work
had been restricted to a subset of the
groups (e.g., A. duplicata, and A.
forsythi), we would have had no reason
to be so cautious. This study thus
illustrates the importance of examining
several groups of related taxa before
trying to reconcile the evolutionary
history of a group with events in the
geological past. Failure to do so can
lead to the phylogeographic equivalent
of adaptationist ‘just-so’ stories.
14
For more information on this study see: Spencer, H.G., Brook, F.J., &
Kennedy, M. 2006. Phylogeography of
Kauri snails and their Allies from
Northland, New Zealand (Mollusca:
Gastropoda: Rhytididae: Paryphantinae).
Molecular Phylogenetics and Evolution,
38, 835-842.
Taxonomy and classification
Our results suggest that the current
taxonomy and classification of these
taxa requires some revision. From our
results you might argue that some
populations of P. busbyi may be better
described as P. watti (or perhaps that
there should be a third Paryphanta
species). You would most likely also
argue that A. dunniae should include A.
tarangaensis, A. pycrofti and A. sp.
“Motukokako”, whereas you might
argue that the A. forsythi we are calling
A. sp. “Aupouri” at the moment are
different enough to warrant specific
status.
Further land snail studies at Otago
We are currently working on several
related studies. The Rhytididae studies
include the one mentioned earlier on
the phylogeny of the Paua Slugs
(Schizoglossa), a study on Rhytida and
Wainuia and a study that combines all
the others and looks at the phylogeny
of New Zealand rhytidids as a whole. A
similar study looks at another group of
snails, the Charopidae. The charopid
study is in its infancy, but will
investigate the phylogeography of
Allodiscus dimorphus and its relatives –
a group that shares large parts of its
distribution with our paryphantine study
– thus allowing us to further investigate
the phylogeographic patterns of
landsnails in Northland.
Recent Publications Baroni, M., Semple, C., and Steel, M. (2006). Hybrids in real time. Systematic Biology 44(1): 46-56: 2006. Chan, C., Ballantyne, K.N., Lambert, D.M. and Chambers, G.K. (2005). Characterization of variable microsatellite loci in Forbes’ parakeet (Cyanoramphus forbesi) and their use in other parrots. Conservation Genetics 6: 651-654. Chan, Z.S.H., Kasabov, N. and Collins, L. (2006). A two-stage methodology for gene regulatory network extraction from time-course gene expression data. Expert Systems with Applications 30:59-63. Chan, Z.S.H., Kasabov, N., and Collins, L. (2005). A hybrid genetic algorithm and expectation maximization method for global gene trajectory clustering. J Bioinf & Comp Biol 3:1227-1242. Chapple, D.G. (2005). Life history and reproductive ecology of White’s skink, Egernia whitii. Australian Journal of Zoology 53: 353-360. Chor, B., Hendy, M.D. and Snir, S. (2006). Maximum Likelihood Jukes-Cantor Triplets: Analytic Solutions, Molecular Biology and Evolution, 23: 626-632 Clarke, A.C., Burkenshaw, M., McLenachan, P.A., Erickson, D. and Penny, D. (2006). Reconstructing the origins and dispersal of the Polynesian bottle gourd (Lagenaria siceraria). Molecular Biology and Evolution 23: 893-900. Collins, L.J. and Penny, D. (2006). Investigating the intron recognition mechanism in eukaryotes. Molecular Biology and Evolution. 23: 901-910. Martyn Kennedy
Research Fellow with Hamish Spencer, University of Otago [email protected]
Donald, K.M., Kennedy, M., and Spencer, H.G. (2005). Cladogenesis as the result of long-distance rafting events in South Pacific topshells (Gastropoda, Trochidae). Evolution 59(8): 1701–1711 Donald, K.M., Kennedy, M. and Spencer, H.G. (2005). The phylogeny and taxonomy of austral monodontine topshells (Mollusca: Gastropoda: Trochidae), inferred from DNA sequences. Mol Phylo Evol 37: 474-483. Duffield, S.J., Winder, L. and Chapple, D.G. (2005). Calibration of sampling techniques and determination of sample size for the estimation of egg and larval populations of Helicoverpa spp. (Lepidoptera: Noctuidae) on irrigated soybean. Australian Journal of Entomology 44: 293-298.
15
Erickson, D.L., Smith, B.D., Clarke, A.C., Sandweiss, D.H. and Tuross, N. (2005). An Asian origin for a 10,000-year-old domesticated plant in the Americas. Proceedings of the National Academy of Science, USA 102(51): 18315-18320. Gartrell, B. and Hare, K.M. (2005). Mycotic dermatitis with digital gangrene and osteomyelitis, and protozoal intestinal parasitism in Marlborough green geckos (Naultinus manukanus). New Zealand Veterinary Journal 53(5): 363-367 Gluckman, P.D., Hanson, M.A., and Spencer, H.G. (2005). Predictive adaptive responses and human evolution. Trends in Ecology and Evolution 20: 527-533. Goremykin, V.V., Holland,B., Hirsch-Ernst, K.I. and Hellwig, F.H. (2005). Analysis of Acorus calamus chloroplast genome and its phylogenetic implications. Molecular Biology and Evolution 22: 1813-1822. Hare, K.M. and Cree, A. (2005). Natural history of Hoplodactylus stephensi (Reptilia: Gekkonidae) on Stephens Island, Cook Strait, New Zealand. New Zealand Journal of Ecology 29(1): 137-142. Hare, K.M., Miller, J.H., Clark, A.G. and Daugherty, C.H. (2005). Muscle lactate dehydrogenase is not cold-adapted in nocturnal lizards from cool-temperate habitats. Comparative Biochemistry and Physiology, Part B 142(4): 438-444. Hendy, M.D. (2005). Hadamard conjugation: an analytic tool for phylogenetics. Chapter 6, pp 143-177, In Mathematics of Evolution and Phylogeny (O.Gascuel ed), Oxford University Press. Hogg, I.D., Stevens, M.I., Schnabel, K.E. and Chapman, M.A. (2006). Deeply divergent lineages of the widespread New Zealand amphipod Paracalliope fluviatilis revealed using allozyme and mitochondrial DNA analyses. Freshwater Biology 51: 236-248. Holland, B. and Schmid, J. (2005). Selecting representative model strains. BMC Microbiology 5: 26. Hörandl, E., Paun, O., Johansson, J.T., Lehnebach, C., Armstrong, T., Chen, L. and Lockhart, P.J. (2005). Phylogenetic relationships and evolutionary traits in Ranunculus s.l. (Ranunculaceae) inferred from ITS sequence analysis Mol Phyl Evol 36: 305-327. Huber, K., Moulton, V. and Steel, M. (2005). Four characters suffice to convexly define a phylogenetic tree. SIAM Journal on Discrete Mathematics 18(4): 835--843.
Huson, D., Kloepper, T., Lockhart, P.J. and Steel, M.A. (2005). Reconstruction of Reticulate Networks from Gene Trees In Proceedings of the ninth international conference in computational molecular biology (RECOMB): 233-249. Jeffares, D.C., Mourier, T and Penny, D. (2006). The biology of intron gain and loss. TRENDS in Genetics 22 (1): 16-22 Johnson, K.P., Kennedy, M. and McCracken, K.G. (2006). Reinterpreting the Origins of Flamingo Lice: Cospeciation or Host-Switching? Biology Letters 2: 275-278. Kennedy, M., Holland, B.R., Gray, R.D. and Spencer, H.G. (2005). Untangling Long Branches: Identifying Conflicting Phylogenetic Signals a priori using Spectral Analysis, Neighbor-Net, and Consensus Networks. Systematic Biology 54:620-633. Kurland, C.G., Collins, L.J., and Penny, D. (2006). Genomics and the Irreducible Nature of Eukaryote Cells. Science 312: 1011-1014. Larson, G., Dobney, K., Albarella, U., Fang, M., Matisoo-Smith, E., Robins, J., Lowden, S., Finlayson, H., Brand, T., Willerslev, E., Rowley-Conwy, P., Andersson L. and Cooper, A. (2005). Worldwide phylogeography of wild boar reveals multiple centres of pig domestication. Science 307:1618-1621. Larson, G., Dobney, K., Albarella, U., Matisoo-Smith, E., Robins, J., Lowden, S., Rowley-Conwy, P., Andersson, L. and Cooper, A. (2005). Response to Domesticated Pigs in Eastern Indonesia. Science 309:381. Lockhart, P.J., Novis, P., Milligan, B.G., Riden, J., Rambaut, A. and Larkum, A.W.D. (2005) Heterotachy and Tree Building: A Case Study with Plastids and Eubacteria. Mol Biol Evol 23: 40-45. Lockhart, P.J. and Penny, D. (2005). The place of Amborella in the radiation of angiosperms. Trends Plant Sci.10: 201-202. Lockhart, P. and Steel, M. (2005). A tale of two processes. Systematic Biology, 54(6): 948-951. McCallum, J., Clarke, A., Pither-Joyce, M., Shaw, M., Butler, R., Brash, D., Scheffer, J., Sims, I., van Heusden, S., Shigyo, M. and Havey, M. J. (2006). Genetic mapping of a major gene affecting onion bulb fructan content. Theoretical and Applied Genetics 112(5): 958-967. McGaughran, A., Hogg, I.D., Stevens, M.I., Chadderton, W.L. and Winterbourn, M.J. (2006). Genetic divergence of three freshwater isopod species from southern New Zealand. Journal of Biogeography 33: 23-30.
Matisoo-Smith, E. (2005). The Rat Path - Tracing Polynesian migration through rat DNA. Wild California - The magazine of the California Academy of Science. 58(2):16-19. Matisoo-Smith, E., Roberts, K., Welikala, N., Tannock, G., Chester, P., Feek, D. and Flenley, J. (2005). DNA and pollen from the same Lake Core from New Zealand. Pp. 15-28 In C.M. Stevenson, J. M. Ramírez Aliaga, F.J. Morin, and N. Barbacci (eds) The Reñaca Papers. VI International Conference on Easter Island and the Pacific/VI Congreso internacional sobre Rapa Nui y el Pacífico. The Easter Island Foundation, Los Osos. ISBN 1-880636-08-5 Miller, H.C., Belov, K. and Daugherty, C.H. (2005). Characterisation of MHC class II genes from an ancient reptile lineage, Sphenodon (tuatara). Immunogenetics 57: 883-891. Morgan-Richards, M. (2005). Chromosome rearrangements are not accompanied by expected genome size change in the tree weta Hemideina thoracica (Orthoptera, Anostostomatidae). Journal of Orthoptera Research, 14(2): 143-148. Ovidiu, P., Lehnebach, C., Johansson, J.T., Lockhart, P.J. and Hörandl, E. (2005). Phylogenetic relationships and biogeography of Ranunculus and allied genera (Ranunculaceae) in the Mediterranean region and in the European Alpine System. Taxon 54: 911-930. Penny, D. (2005). An interpretive review of the origin of life research. Biology and Philosophy 20:633–671 Perrie, L.R., Shepherd. L.D. and Brownsey, P.J. (2005). Asplenium xlucrosum nothosp. Nov.: a sterile hybrid widely and erroneously cultivated as “Aspelium bulbiferum”. Plant Syst Evol 250:243-257. Phillips, M.J. (2006). Sympathy for the Devil. Nature 440. Phillips, M.J., McLenachan, P.A., Down, C., Gibb, G.C. and Penny, D. (2006). Combined nuclear and mitochondrial protein -coding DNA sequences resolve the interrelations of the major Australasian marsupial radiations. Systematic Biology 55: 122-137. Robins, J.H., Ross, H.A., Allen, M.S. and Matisoo-Smith, E.M. (2006). Sus bucculentus revisited. Nature 440. Semple, C. and Steel, M. (2006). Unicyclic networks: compatibility and enumeration. IEEE/ACM Transactions on Computational Biology and Bioinformatics 3(1), 84-91.
16
Shepherd, L.D. and Lambert, D.M. (2006). Nuclear microsatellite DNA markers for New Zealand kiwi (Apteryx spp.). Molecular Ecology Notes 6: 227-229.
Stevens, M.I. and Hogg, I.D. (2006). Molecular ecology of Antarctic terrestrial invertebrates and microbes. Chapter 9 in: Trends in Antarctic Terrestrial and Limnetic Ecosystems: Antarctica as a global indicator. Eds. A. Huiskes, P. Convey and D. Bergstrom. ISBN 1-4020-5276-6. Springer, Dordrecht, The Netherlands.
Contact Us Allan Wilson Centre for Molecular Ecology and Evolution Shepherd, L.D. and Lambert, D.M. (2005).
Mutational drive in penguin microsatellite DNA. Journal of Heredity 96(5): 566-571.
Host Institution Massey University, Private Bag 11 222, Shepherd, L.D., Millar, C.D., Ballard, G.,
Ainley, D.G., Wilson, P.R., Haynes, G.D., Baroni, C. and Lambert D.M. (2005). Microevolution and mega-icebergs in the Antarctic. Proceedings of the National Academy of Sciences USA 102: 16717-16722.
Stevens, M.I., Hogg, I.D. and Pilditch, C.A. (2006). Evidence for female-biased juvenile dispersal in corophiid amphipods from a New Zealand estuary. Journal of Experimental Marine Biology and Ecology 331: 9-20.
Palmerston North, New Zealand Phone: 64 6 350 5448 Fax: 64 6 350 5626
Partner Institutions Winkworth, R.C., Wagstaff, S.J., Glenny, D. and Lockhart P.J. (2005). Evolution of the New Zealand alpine flora: origins, diversification and dispersal. Org. Divers. Evol. 5: 237-247
The University of Otago, P. O. Box 56 Dunedin, New Zealand
Slack, K.E., Jones, C.M., Ando, T., Harrison, G.L., Fordyce, E., Arnason, U. and Penny, D. (2006). Early penguin fossils, plus mitochondrial genomes, calibrate avian evolution. Molecular Biology and Evolution 23: 1144-1155
The University of Auckland, Private Bag 92019
Zauner, S., Lockhart, P.J., Stoebe-Maier, B., Gilson, P., McFadden, G.I. and Maier, U.G. (2006). Differential Gene Transfers and Gene Duplications in Primary and Secondary Endosymbioses. BioMed Central 6: 38.
Auckland, New Zealand Victoria University of Wellington, P. O. Box 600 Spencer, H.G., Brook, F.J. and Kennedy, M.
(2006). Phylogeography of Kauri Snails and their Allies from Northland, New Zealand (Mollusca: Gastropoda: Rhytididae: Paryphantinae). Molecular Phylogenetics and Evolution 38: 835-842.
Wellington, New Zealand Canterbury University, Private Bag 4800 Christchurch, New Zealand Spencer, H.G. and Feldman, M.W. (2005). Adaptive dynamics, game theory and evolutionary population genetics. Journal of Evolutionary Biology 18: 1191-1193.
Production Editor: Steel, M. (2005). Phylogenetic diversity and the greedy algorithm. Systematic Biology 54(4): 527-529.
Susan Wright Assistant Editor, Design and Layout: Nathalie Loussert Printed by: Steel, M. (2005). Should phylogenetic
models be trying to `fit an elephant'? Trends in Genetics 21(6): 307-309.
Massey University Printery Newsletter banner design and photo credits: Nathalie Loussert Steel, M. and Hein, J. (2006). Reconstructing pedigrees: a combinatorial perspective. Journal of Theoretical Biology 240(3): 360-367.
© The Allan Wilson Centre 2006. The Allan Wilson Centre Newsletter is available on request. Email Susan Wright at [email protected]
Steel, M. and Pickett, K.M. (2006). On the
impossibility of uniform priors on clade size. Molecular Phylogenetics and Evolution 39(2): 585-586.
Visit the Allan Wilson Centre website at: http://awcmee.massey.ac.nz
Stevens, M.I., Greenslade, P., Hogg, I.D. and Sunnucks, P. (2006). Examining Southern Hemisphere springtails: could any have survived glaciation of Antarctica? Molecular Biology and Evolution 23: 874-882.
Any information in this newsletter may be reused provided The Allan Wilson Centre is acknowledged as the source of the information.
(Recent publications can be viewed by visiting the Allan Wilson Centre: http://awcmee.massey.ac.nz/publications.htm)