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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
Th T he e A A l l l l a a n n W W i i l l s s o o n n C C e e n n t t r r e e N N e e w w s s l l e e t t t t e e r r
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).
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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 Stevens Postdoc,
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.
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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
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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 Lawrence PhD 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?
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
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.
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.
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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).
Mutational drive in penguin microsatellite DNA. Journal of Heredity 96(5): 566-571.
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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.
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