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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).

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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 A.C.Clarke@massey.ac.nz

English to Māori translation by Māori Language Services, Māori Language Commission – Te Taura Whiri i te Reo Māori.

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‘Giant’ Collembola of New Zealand: The Largest Springtails in the World!

What do Collembola do?

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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 m.i.stevens@massey.ac.nz

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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 H.Lawrence@massey.ac.nz

Phylogeography of Carnivorous Land Snails (Family Rhytididae)

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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.

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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?

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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.

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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 martyn.kennedy@stonebow.otago.ac.nz

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.

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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.

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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).

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