The Allan Wilson Centre Newsletter

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Issue Number 4 (September 2006) Origins and Dispersal of the Ngā Orokohanga me ngā Polynesian Bottle Gourd Tuaritanga o te Hue o Te

Moana nui a Kiwa

Our research group at the Allan Wilson Inside this issue Kua kitea e tō mātou rōpū i te Allan Centre have discovered the bottle Wilson Centre, ko te Hue (he Bottle Origins and Dispersal of the gourd (or hue in Māori) grown in Gourd i te reo Ingarihi) e tipu ana ki Te Polynesia originated in both Asia and Polynesian Bottle Gourd ...... 1 Moana nui a Kiwa, i taketake mai i Āhia the Americas. The bottle gourd, which me ngā motu o Amerika. Ko te hue Ngā Orokohanga me ngā is closely related to the pumpkin, is one tētahi o ngā huawhenua maha i kawea of the many crops that Polynesians Tuaritanga o te Hue o Te Moana mai e ngā tāngata o Te Moana nui ā took with them as they settled the nui a Kiwa ...... 1 Kiwa i a rātou e noho haere ana i ngā islands of the Pacific, including moutere o te Moana nui ā Kiwa, tae Aotearoa New Zealand. ‘Giant’ Collembola of New noa ki Aotearoa. He whanaunga tata te

Zealand: The Largest Springtails hue ki te paukena. Anthropologists had previously in the World!...... 4 suggested the bottle gourd had come I ngā rā ki muri, i kī ngā tohunga

from South America along with the tikanga tangata i taketake kē mai te Tuatara Assisting with Education sweet potato (kumara), but our hue i Amerika ki te Tonga i te taha o te Outreach ...... 7 research shows there is also a kūmara, heoi kua kitea i roto i tō mātou

significant genetic contribution from rangahau, te kaha uru o ngā momo Celebration of Te Kopinga, First Asia, and that Polynesian bottle gourds whakaheke mai i Āhia ā, ko te mea kē, Marae of the Moriori ...... 8 are in fact hybrids between gourds from he kākano whakauru kē nō ngā motu e both of these continents. rua nei. Phylogeography of He tino maha ngā kākano hue i Carnivorous Land Snails We collected a large number of bottle gourds seeds from Asia and the kohikohia e mātou i Āhia me ngā wāhi (Family Rhytididae) ...... 10 Americas, as well as eight Māori bottle o Amerika whānui, tae noa ki ngā hue e

gourds from New Zealand. waru o Aotearoa. Recent Publications ...... 14

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). 1

The Māori gourds were obtained from marae and heritage I tīkina mai ngā hue Māori i ngā marae me ētahi kamupene

seed companies, and are thought to be derived from true pupuri ā-tikanga i ngā kākano ā, ko te whakaaro, i ahu mai Māori bottle gourds grown in pre-European New Zealand. ēnei i ngā hue a te Māori i whakatipuria i mua i te taenga mai o te Pākehā. This collection was used to develop DNA markers that could be used to trace the gourd’s origins. We used DNA I whakamahia tēnei kohikohinga hei hanga tohu pītau-ira fingerprinting, similar to that used to identify humans, to (DNA) hei whakataki i te takenga mai o te hue. I whakamahia locate regions of the gourd genome that are variable. Just e mātou te tapukara pītau ira (DNA) rite ana ki tērā e as in humans, individual bottle gourds share nearly whakamahia ana ki te tautuhi i te tangata, hei rapu i ngā wāhi identical DNA – probably more than 99% – so the DNA tipu ai te hue whai tāupe. He tino ōrite katoa nei ngā pītau-ira fingerprinting is used to identify the less than 1% of the o ia hue, pērā anō ki te tangata – te āhua nei nui atu i te 99 DNA that makes each bottle gourd different. These ōrau – nā reira ka whakamahia te tapukara pītau-ira hei variable DNA fragments could then be used as DNA tautuhi i te toenga o te 1 ōrau o te pītau-ira, e rerekē ai tēnā markers to trace the origins of the Polynesian bottle gourd. hue ki tēnā hue. Ka taea ēnei maramara pītau-ira tāupe te whakamahi hei kai tohu pītau-ira, hei whakataki hoki i te The DNA markers showed that Asian gourds are all of one takenga mai o te hue o Te Moana nui a Kiwa. type, American gourds are all of another type, and that Polynesian gourds are a mixture of both. This opens a I whakaaturia mai e ngā kaitohutohu pītau-ira he momo kotahi number of possibilities for the dispersal of this species. 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. 2 (Photo credit: Archives New Zealand and the Alexander Turnbull Library, Wellington).

Bottle gourds could have been brought from Asia with the Tērā pea i haria mai te hue i Āhia e ngā tūpuna o Te Moana ancestors of Polynesians when they moved out of South nui ā Kiwa i te wā i puta ai rātou i Āhia ki te tonga, e 5,000 tau East Asia 5,000 years ago, or perhaps with later migrants ki muri. Tērā pea i haria mai ngā hue Amerikana ki Te Moana from Asia. The American gourds could have been nui ā Kiwa i te taha o te kūmara: Inā hoki, tērā tētahi kōrero, i introduced to Polynesia with the kumara. Polynesian haere ngā kaiwhakatere waka o Te Moana nui ā Kiwa mai i voyagers are thought to have sailed from Easter Island to Rapanui ki Amerika ki te Tonga, he āhua 1,000 tau ki muri, i South America about 1,000 years ago and collected the reira kohikohi ai i te kumara i mua i tōna hokinga ki te Moana- kumara before sailing back to Polynesia. The bottle gourd nui-Kiwa. He tino māngi te hue, nā reira ko wai ka mōhio tērā is also very buoyant, so we cannot rule out that a gourd pea i māunu kē mai i Āhia, mai i ngā wāhi o Amerika rānei, ki floated from Asia or the Americas to Polynesia, where it Te Moana nui ā Kiwa. I reira ka kohia mai te ākau ka was picked up from a beach and propagated from the whakamakuru ai ngā kākano o roto i te hue. seeds which are stored inside the fruit. Ko te hue tētahi o ngā momo huawhenua tino hira i mua i te The bottle gourd was one of the most important crop taenga o ngāi Pākehā ki Te Moana nui ā Kiwa. I kainga ngā species in pre-European Polynesia. In New Zealand young hue iti (pērā ki te zucchini) engari i tino whakamahia ngā hue i bottle gourds were eaten (like zucchini), but were mainly te wā kua hua, kua maroke hoki. I hākarohia ēnei hue mārō used when dry and mature. These hard-shelled bottle nei ana, kātahi ka whakamahia hei oko kawe wai, hei ipu, hei gourds were hollowed out and used primarily as water- kūmete kai rānei (ka huahuatia ngā tītī me ngā tūi) hei taonga carrying vessels, containers for food (muttonbirds and tui pūoro, hei tīheru mō te waka. E whakatipua tonu ana te hue were stored in their own fat), musical instruments and a te Māori, i Aotearoa nei, heoi hei whakapaipai noa iho, canoe bailers. Māori bottle gourds are still grown in New (pērā ki tā te pikitia nei) i te nuinga o te wā, me te whakaora Zealand today, but mostly for ornamental purposes (such tonu i tēnei āhuatanga o Te Moana nui ā Kiwa me te ao o te as the one pictured) and to preserve this important part of Māori. Polynesian and Māori life.

Acknowledgement: NGĀ MIHI We are grateful to Mr Jonathan Ngā mihi ki a Jonathan Procter me Rangitāne Procter and Rangitaane O o Manawatū mō tā rātou tautoko i ngā mahi ira Manawatu for supporting the tangata i whakahaerehia e pā ana ki te hue. genetic work undertaken on the bottle gourd.

Andrew Clarke English to Māori translation by PhD student Māori Language Services, Massey University Māori Language Commission – [email protected] Figure 3: Ornamental bottle gourd carved with Te Taura Whiri i te Reo Māori.

modern Māori design. (Photo credit: Andrew Clarke).

‘Giant’ Collembola of New tasmaniae, Acanthanura dendyi, worms (Peripatus), but the Zealand: The Largest Womersleymeria bicornis), and New Uchidanurinae are currently only Zealand (Holacanthella spinosa, H. considered to be of extreme Springtails in the World! paucispinosa, H. brevispinosa, H. conservation status in Australia. They

duospinosa, H. laterospinosa). are likely to be a particularly important What do Collembola do? part of New Zealand’s saproxylic fauna Collembola (springtails) are an ancient These species are particularly as springtails have been shown to be (>412MYA) and highly successful class remarkable in that some are the largest key agents in controlling the dynamics of hexapod dating back to at least the springtails recorded world-wide (up to of soil microorganisms (bacteria, fungi Devonian (Rhyniella praecursor) or 17 mm long for the New Zealand and algae), and thus play a crucial role Upper Silurian. Although predominantly species), and most sport coloured in defining the composition of the soil and litter dwellers, they also occur digitations (spine-like projections) on saproxylic community. in a wide range of habitats such as on their dorsal and lateral surfaces (Fig. vegetation, under rocks, in logs (Fig. 1), 2), and are saproxylic (live within in tree canopies, in caves, in the marine decomposing logs). littoral zone, and in freshwater systems. Saproxylic Communities As detritivores, springtails are an Saproxylic communities drive important group in nutrient cycling and nutrient cycling and nutrient are beneficial organisms as very few uptake by plants in forests. This species feed on live plant material. The action returns nutrients locked up ecology and widespread nature of in dead wood to the ecosystem springtails suggest that they warrant where they support large and more attention from biologists. diverse invertebrate populations Worldwide, over 7000 species in 581 and enrich the soil to enhance genera have been described and are growth and regeneration. A large found throughout the world including proportion of the New Zealand the Arctic and Antarctic regions. endemic plants and animals Figure 1: A typical habitat within a South Island (Arthurs

considered to be of conservation Pass) beech forest. (Photo credit: Mark Stevens). The ‘Giants’ importance are adapted to native The most spectacular and largest forests and saproxylic springtails form the subfamily communities are an important part Uchidanurinae Salmon, 1964. The of these ecosystems. As well as Uchidanurinae currently consists of enriching forest soils, saproxylic eight genera and 15 species all of organisms (which include, for which are endemic to their respective example, earthworms, myriapods, localities—China (Assamanura fungi, beetles and spiders) provide besucheti), Indo-China (Denisimeria important food sources (directly Figure 2: Collembola (Holacanthella duospinosa.) collected caudata, D. longilobata, D. martyni), and indirectly) for a number of from Ohakune. Known to reach 17mm in length, which Micronesia/Polynesia (Uchidanura New Zealand’s most treasured makes it the largest known springtail in the world. bellingeri, U. esakii), New Caledonia and threatened species including (Photo credit: Rod Morris). (Caledonimeria mirabilis), eastern Kiwi, rhytidid snails (including the Australia/Tasmania (Megalanura Powelliphanta), robins and velvet 4

What are we doing? records determining what effect New Caledonian species using Despite the overwhelming ecological disturbances have had on these unique mitochondrial and nuclear DNA. importance of New Zealand’s and important springtails has been an (4) Examine the phylogeographic Uchidanurinae they have not been the arduous task. patterns for the three widespread New subject of scientific interest since their Zealand species (H. brevispinosa, H. original descriptions between 1899 and Our work aims to: paucispinosa, H. duospinosa) using 1944. In recent times the forests they (1) Provide a detailed examination of mitochondrial and anonymous nuclear inhabit have undergone large scale the distribution of all five Holacanthella markers. fragmentation following first Polynesian, species throughout New Zealand. then European settlement, and (2) Develop an updated key to their Distribution of all five Holacanthella subsequent infestation by introduced identification (available online: species throughout New Zealand pests. Future scientific and/or http://awcmee.massey.ac.nz/people/ms Throughout New Zealand the density of conservation effort requires a greater tevens/NZ.htm) Holacanthella individuals found at any understanding of their distribution, but (3) Examine phylogenetic relationships particular site was highest in beech with only a total of eleven historical for the New Zealand, Australian and 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 other mixed forest types. Several sites Holacanthella are an under-studied possessed more than one species, for group that are likely to be an important example locations in southland, part of New Zealand’s forgotten Fiordland, Arthurs Pass, Lewis Pass, invertebrate biota. At present the Mt Arthur Tableland, Wellington, and Department of Conservation Ohakune. The distribution of the two Invertebrate recovery plan makes no species H. brevispinosa and H. mention of any of New Zealand’s paucispinosa was almost completely springtail species. Collembola are overlapping (sympatric) extending typically considered too small and too from Stewart Island, throughout South numerous to be considered in need of Island, and extended north to the conservation. However, this is not Figure 4: Holacanthella paucispinosa collected from Rahu Central Plateau of North Island. Saddle, South Island. (Photo credit: Rod Morris). always the case: a reserve in Finding both species in or under a Tasmania (Springtail Reserve) has single log occurred on several been dedicated solely for a species of occasions. Holacanthella spinosa is Conserving forgotten species Collembola, Tasphorura vesiculata; the only other South Island species. At The loss of habitat emphasises human species of Uchidanurinae were listed by Mt Ruapehu, on the Central Plateau, impacts which is currently likely to be the IUCN in the Red Data book in 1994; the three species were found together the greatest threat to this group. Most and the removal of dead wood is listed with another species, H. duospinosa, importantly, available dead wood on the as a threatening process in NSW, and this species extends north to forest floor is a requirement of these Australia. The likely ecological Northland (including Kawau I., Little saproxylic communities. ‘Natural’ importance and the vulnerability of Barrier I., Great Barrier I.). The fifth forests (unmanaged) currently support Holacanthella means they should form remaining species, H. laterospinosa, is large populations of Holacathella. Most a part of future conservation plans. only known from Cuvier Island off the notable are beech forests that have not Coromandel Peninsula (North Island). undergone extensive logging, such as Understanding of their distribution and in southland, Abel Tasman National genetic diversity will aid in determining With a recent summer student (David Park (Mt Arthur tableland), and the vulnerable/rare species and regions. Winter) and numerous field helpers we Tongariro National Park (Central Our objective is to understand more have extended the known distribution of Plateau), all support dense, species fully ‘the small things that run the world’ all five New Zealand endemic rich populations. However, most of New and the processes that have shaped Holacathella species. The historic Zealand’s remnant forests are broken New Zealand’s biodiversity. (MONZ) and new records highlight the into small fragments. In total there are importance of maintaining old growth around 120,000 such fragments with an For further reading: forests in the west coast and northern average size of 53.9 Ha. Collembola Collins Field Guide to New Zealand Wildlife. South Island, central North Island, and are known to be highly sensitive to By Terence Lindsey and Rod Morris. Page 187. ISBN 1-86950-300-7 Cuvier Island to adequately preserve forest practices and their low dispersal these species. Molecular and capacity makes recolonisation of morphological studies are now disturbed (and regenerating) sites more underway to further examine the difficult, particularly if these are Mark Stevens Postdoc, intraspecific (within species) fragmented. The preservation of the Massey University, morphological variability that we ‘natural’ characteristics of these Palmerston North [email protected] observed across the ranges for these habitats and their original species species. composition appears essential. 6

Tuatara Assisting with Greymouth. Presentations were given Education Outreach at 57 primary schools, 12 secondary schools and 9 public venues. Each

school group consisted of 50 students “Tuatara: a Taonga for the People of and several teachers (limited in size for New Zealand”, a joint project between the welfare of the tuatara): public Victoria University of Wellington, Te groups ranged in size up to 100. Atiawa iwi, The Allan Wilson Centre, and Approximately 3500 members of the the San Diego Zoo was successfully New Zealand public participated in the completed in December 2005. Funded programme in total. The presentations by the Royal Society of New Zealand’s were enthusiastically received, and the Science and Technology Promotion opportunity to meet and touch a live Fund, this project took conservation tuatara had real impact. Substantial education outreach about tuatara to feedback about the educational value schools around New Zealand. An of the presentation was received, with additional goal of the project was to extremely positive comments such as provide training to iwi members in tuatara “a fabulous presentation and one which biology, research and conservation the students will remember always”. education. Media interest was also high, with 19

newspaper articles reporting the Training began when two iwi project’s school and public representatives attended a Conservation presentations. Education Workshop hosted by the San

Diego Zoo in October 2004. The next The project has enabled iwi presenters step was for them to participate in a to develop knowledge and skills that Victoria University research field trip to will assist them in developing further North Brother Island in March 2005. conservation education outreach Here they gained first-hand knowledge of programmes within their own rohe. An tuatara biology and behaviour, and additional outcome has been the learned techniques in scientific research. Project on Tuatara conservation throughout schools in positive example set by these young iwi New Zealand. (Photo credits: Sue Keall). teachers to their peers, as to what can In April there was a week long visit to be achieved with further training in conservation of native biodiversity. The Victoria University in which the two science and conservation of our taonga. presentation concluded with a live teachers built on their knowledge of tuatara being shown to attendees on an tuatara biology, the results of scientific individual basis, and in most cases research and how it is being applied to touching the tuatara was encouraged. tuatara conservation. Several

Wellington primary schools were visited Once training was complete, the project so that trial presentations could be visited schools in five centres around given. A 30 minute narrated New Zealand during 2005 – Sue Keall Powerpoint presentation conveyed how Technical Officer Blenheim/Picton, New Plymouth, Victoria University of Wellington science and technology play an Whakatane, Whangarei and essential role in supporting the 7

Celebration of Te Kopinga, was thought to be extinct as were the that by air it looks like an albatross in First Marae of the Moriori Moriori people themselves, but the flight. The albatross is also a taonga Moriori and the Taiko did survive. The species to the Moriori. “Hokomenetai” is

Taiko was rediscovered on the night of the name of the wharenui, a house of Last year I was given a very special New Year’s day 1978, by David peace. It is in a pentagonal shape opportunity, to attend the opening Crockett. Mainstream New Zealand emulating the rocks in the basalt celebration of the first Moriori marae on became aware of Moriori survival when columns found on the island. Rekohu/ Chatham Island. I was invited a documentary was filmed in 1980. because of my involvement with the After this, the Moriori people Around 1000 people were present at Moriori (indigenous people) through my commissioned a book by Michael King the opening celebrations. A dawn research. about their history, language and ceremony began the day, very

culture. A Waitangi Tribunal claim in appropriate since the Chathams is the 2001 brought official recognition of their first place to see the sun. At lunchtime unique status. The opening of the first the official whakamaurahiri (welcoming) Moriori marae was a great achievement began. It was slightly different from a in the reaffirmation of culture and the Maori powhiri. Moriori Kuia called the joining together of Moriori people. maurahiri (manuhiri/visitors) on to the marae while the Ratana church band My PhD is on the conservation genetics Te Kopinga is the first Moriori marae led us. We then entered the wharenui of the Chatham Island Taiko, called because instead of marae, Moriori used for the hau-rongo (speeches). There Tchaik by Moriori. This petrel is New to meet in groves of Kopi (Karaka) was no wero (challenge) because the Zealand’s most endangered seabird. In trees. The new marae looks over Te Moriori live under Nunuku’s law, a the past it was an important food Awapatiki, the mouth of the lagoon, covenant of peace. Karakii were said source for some Moriori imi (iwi/tribes). where all imi met in the past. The as Ka Pou o Rangitokona (the central The birding practice was highly building complex of the wharenui (main post) was blessed. Moriori Rangata ritualised and involved special karakii house), kitchen, dining hall, and Matua (Kaumatua/elders) and leaders (karakia/prayers). Conservation was administration blocks, was designed so spoke, some in Moriori, some in Maori, 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.

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 Figure 1. View of Te Awapatiki, mouth of Te Whanga (lagoon), ancient meeting place 8 of the Moriori. (Photo credit: Hayley Lawrence).

After the ceremony was kai time. The and made new ones. The organisers feast was amazing and included a genuinely thanked me for coming, bounty of kaimoana (seafood). People which surprised me because I felt exclaimed in delight at the size of the honoured just to be asked. I think that koura (crayfish) that the Chathams is they appreciated my attendance as renowned for. Other delicacies included demonstrating that our relationship is akoako (titi/muttonbird), hangi, and important to me. I distributed weka (a bird which can only legally be information to interested people and eaten on the island). After the feed, displayed a poster. It included a Moana and the tribe (formerly Moana request for information regarding and the Moahunters) entertained us. traditional ecological knowledge, but

After that many people headed off to also included an invitation for people to and some in English. Moriori rongo the legendary Hotel Chathams, which contact me if they would like to know (waiata/songs) were sung after each has its own Island Gold beer, White more about my work (reciprocity is speaker. Speeches were made by Pointer vodka, and Blind Jims bourbon. important). (The poster is also available invited guests, including Kaumatua (I was warned about the bourbon!) on the AWC website at: from Maori iwi from across Aotearoa http://awcmee.massey.ac.nz/project_H and Te Wai Pounamu (NZ). Michael The trip was fun, but also useful for my Lawrence.htm) King’s son spoke in his honour, project. Imi/Iwi consultation is essential followed by Helen Clarke. At the for a few aspects of my work, including I also visited other people I know in the conclusion of the speeches, Moriori cloning and bone collecting. I believe it Taiko Trust and Department of children, and Maori and Pakeha is important to establish good Conservation. They have helped me children from the island renewed the relationships with imi/iwi, especially greatly with my project, especially covenant of peace. During this moving when the species you are working on is logistically, for which I am very thankful. ceremony, Moana and the tribe (the a taonga to them. The relationship can Getting to know them on a personal band) sang a song specially composed be very rewarding for both sides. level has been great. All in all, my visit for the occasion. During my trip, I reaffirmed old contacts 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 [email protected]

Figure 2. Hokomenetai, the wharenui of Te Kopinga marae. (Photo credits for this page: Hayley Lawrence). 9

Phylogeography of poorly understood, and many Carnivorous Land Snails of the group’s members are of conservation concern (mostly (Family Rhytididae) due to rat predation and

habitat destruction). New Zealand Rhytididae

The New Zealand Rhytididae are a To be able to address issues large group of carnivorous land snails about the conservation status that include the well known of the members of this group, Powelliphanta group. Members of this we first need to be sure what large family can also be physically very taxonomic groups we are large – at up to 90 mm some actually dealing with. One Figure 2: Amborhytida dunniae. (Photo credit: Fred Brook). Powelliphanta are New Zealand’s way in which we can do this is largest land snails. Other members of to compare the results from using Schizoglossa (Paua Slugs, found in the the group, like Paryphanta, may get up molecular markers with the northern half of the North Island). to 75 mm. With spectacular shells, expectations you would have from these snails have a Gondwanan morphology. By utilizing sequence In this initial study we set out to distribution: that is, they are found data we can evaluate any potential investigate the relationships of the taxa throughout New Zealand, Australia, classification problems there may be restricted to Northland and to place New Guinea, and South Africa. due to either conserved morphologies those relationships within a geographic

or rapidly changing morphological context. Thus we focused on characters (or a combination of these). Paryphanta and Amborhytida, the While investigating the classification of genera restricted to, but widespread these snails is a useful and worthwhile within, Northland. The Kauri Snails purpose on its own, these studies can contain two species, Paryphanta busbyi be put into context by investigating the (Figure 1) (found from Awanui to evolutionary history of the groups – Warkworth, which grows up to 75 mm) including looking at their and Paryphanta watti (found in the Far phylogeography. North only, growing up to 60 mm).

Amborhytida contains five nominal The Paryphantinae: the Kauri Snails species, three of which have and Relatives reasonably wide distributions: The first study of New Zealand Amborhytida dunniae (Figure 2) (found Rhytididae that we have completed is Figure 1: Payphanta busbyi. from Awanui to Auckland), Amborhytida one on the Kauri snails and their (Photo credit: A. M. Spurgeon, supplied from forsythi (found from Karikari to north relatives. In this group most of the New Zealand Mollusca website: Kaipara, and historically considered species are restricted to Northland. http://www.mollusca.co.nz/) closest to Amborhytida dunniae), and Within the Paryphantinae there are four Amborhytida duplicata (from in the Far Rhytidids are carnivorous on worms, as genera of large species: Paryphanta North only). well as other snails and slugs. (Kauri Snails, found throughout

Although these are our largest and Northland), Rhytidarex (from the Three perhaps most charismatic land snails, Kings Islands), Amborhytida (found their classification is still relatively throughout Northland), and 10

Questions Answers the samples reached Otago, we Our initial questions included asking, To answer these questions we needed sequenced an ~1 kb fragment of the what are the evolutionary relationships to generate a phylogeny for the group. mitochondrial COI gene for each of among these species? What are the We (meaning our collaborator Fred them. Various methods were used to evolutionary relationships among Brook) collected samples of all the estimate the phylogenetic relationships populations within these species? And species of the four paryphantine genera of the group – all of which gave very to place it all into some wider context, found in Northland. The samples were similar results. The Bayesian tree do these relationships make sense collected from between three and produced is shown in Figure 3. This geographically and geologically? eighteen locations in Northland. After 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.

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]). 11

Firstly, the phylogeny of Paryphanta group. The morphologically divergent Motukokako (Piercy Island) and nearby does not correspond with the current forms restricted to some of the islands Cape Brett peninsula – fitted clearly taxonomy of the genus. The two off the eastern coast of Northland – A. within the genetic variation ascribed to populations of the Far-North endemic, tarangaensis from Taranga (Hen) A. dunniae. This result suggests that P. watti (represented on Figure 4 by Island, A. pycrofti from the Poor Knights populations of each of these island (or orange stars), that we sampled fell Islands and A. sp. “Motukokako” from near island) endemics are very closely within a clade including several 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 Figure 4. Bayesian tree and map of sample locations for the Paryphanta group. 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 Figure 5. Bayesian tree and map of sample locations for the Amborhytida dunniae group. 12

related, possibly as a consequence of Figure 6) were weakly grouped with A. “Aupouri” evolved on what is now evolutionarily recent founder events. duplicata (the orange stars on Figure Mount Camel or Karikari Peninsula, 6), which is endemic to the area which were also separate islands in the The remaining populations of between Cape Maria van Diemen and Pliocene, before spreading south to Amborhytida, originally attributed to A. North Cape at the northern tip of the Herekino. A. forsythi presumably forsythi and A. duplicata, formed a Aupouri Peninsula. Populations of evolved in mainland Northland. group with very strong support and morphologically similar A. forsythi from were considerably divergent from A. elsewhere in Northland between Taipa What does it all mean? dunniae (see Figure 3). Thus, the view (the type locality) and north Kaipara, For the Amborhytida forsythi group of A. forsythi as only subspecifically formed a separate, well supported (Figure 6) we find interesting and distinct from A. dunniae is not tenable, clade (the blue stars on Figure 6). unexpected patterns, with A. duplicata and in fact the two taxa are locally The almost simultaneous evolution of falling within the group. This result microsympatric (e.g., at locations 18 A. duplicata, A. forsythi, and A. sp. raises interesting questions about the and 19; see Figures 5 and 6). “Aupouri”, which we estimate at being morphological characters that have Moreover, the samples originally between 1.9 and 6.6mya, accords with previously been used to determine the identified from shell morphology as A. the inferred former existence of islands relationships within these groups. The forsythi grouped in a most unexpected in the Cape Reinga-North Cape, Mt phylogeographic patterns within the way (Figure 6), falling into two well- Camel and Karikari areas during Amborhytida forsythi group make sense supported non-sister clades, although Pliocene time (1.8–5.3mya). Clearly, A. – what makes less sense is that that the non-sisterhood itself was not well duplicata evolved in the Far North and they in no way resemble the patterns supported. Populations from Mt Camel, remained there, with eastern and within either the Amborhytida dunniae Karikari Peninsula, and hill country western populations subsequently group or the Paryphanta group. north of Herekino Harbour, becoming genetically (but not Because these snails are closely subsequently referred to in our study as conchologically) differentiated over the related, have similar life histories and A. sp. “Aupouri” (the green stars on last 0.9–3.2my. Possibly, A. sp. 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. 13 reformation of Northland, or a A. sp. “Aupouri” at the moment are Recent Publications combination of these and other different enough to warrant specific processes, we cannot tell at this point. status. Baroni, M., Semple, C., and Steel, M. What these results do tell us is that if (2006). Hybrids in real time. Systematic Biology 44(1): 46-56: 2006. we had looked at just one of these Further land snail studies at Otago groups assuming that, because they We are currently working on several Chan, C., Ballantyne, K.N., Lambert, D.M. and Chambers, G.K. (2005). were closely related to one another and related studies. The Rhytididae studies Characterization of variable microsatellite loci in Forbes’ parakeet (Cyanoramphus had similar life histories and lived in the include the one mentioned earlier on forbesi) and their use in other parrots. same areas, we could generalise from the phylogeny of the Paua Slugs Conservation Genetics 6: 651-654. one group to another we would have (Schizoglossa), a study on Rhytida and Chan, Z.S.H., Kasabov, N. and Collins, L. (2006). A two-stage methodology for gene been very wrong. The discordance in Wainuia and a study that combines all regulatory network extraction from time- course gene expression data. Expert the phylogeographic patterns in the the others and looks at the phylogeny Systems with Applications 30:59-63. groups of snails examined here means of New Zealand rhytidids as a whole. A Chan, Z.S.H., Kasabov, N., and Collins, L. that it is difficult to make strong similar study looks at another group of (2005). A hybrid genetic algorithm and expectation maximization method for global inferences about common geological snails, the Charopidae. The charopid gene trajectory clustering. J Bioinf & Comp influences on the evolutionary history of study is in its infancy, but will Biol 3:1227-1242. paryphantines in Northland. If our work investigate the phylogeography of Chapple, D.G. (2005). Life history and reproductive ecology of White’s skink, had been restricted to a subset of the Allodiscus dimorphus and its relatives – Egernia whitii. Australian Journal of Zoology groups (e.g., A. duplicata, and A. a group that shares large parts of its 53: 353-360. forsythi), we would have had no reason distribution with our paryphantine study Chor, B., Hendy, M.D. and Snir, S. (2006). Maximum Likelihood Jukes-Cantor Triplets: to be so cautious. This study thus – thus allowing us to further investigate Analytic Solutions, Molecular Biology and illustrates the importance of examining the phylogeographic patterns of Evolution, 23: 626-632 several groups of related taxa before landsnails in Northland. Clarke, A.C., Burkenshaw, M., McLenachan, P.A., Erickson, D. and Penny, D. (2006). trying to reconcile the evolutionary Reconstructing the origins and dispersal of history of a group with events in the the Polynesian bottle gourd (Lagenaria siceraria). Molecular Biology and Evolution geological past. Failure to do so can 23: 893-900. lead to the phylogeographic equivalent Collins, L.J. and Penny, D. (2006). Investigating the intron recognition of adaptationist ‘just-so’ stories. mechanism in eukaryotes. Molecular Biology Martyn Kennedy and Evolution. 23: 901-910. Research Fellow with Taxonomy and classification Donald, K.M., Kennedy, M., and Spencer, Hamish Spencer, H.G. (2005). Cladogenesis as the result of Our results suggest that the current University of Otago long-distance rafting events in South Pacific [email protected] taxonomy and classification of these topshells (Gastropoda, Trochidae). Evolution 59(8): 1701–1711 taxa requires some revision. From our Donald, K.M., Kennedy, M. and Spencer, results you might argue that some H.G. (2005). The phylogeny and taxonomy populations of P. busbyi may be better of austral monodontine topshells (Mollusca: Gastropoda: Trochidae), inferred from DNA For more information on this study described as P. watti (or perhaps that sequences. Mol Phylo Evol 37: 474-483. see: Spencer, H.G., Brook, F.J., & there should be a third Paryphanta Duffield, S.J., Winder, L. and Chapple, D.G. Kennedy, M. 2006. Phylogeography of species). You would most likely also (2005). Calibration of sampling techniques Kauri snails and their Allies from and determination of sample size for the estimation of egg and larval populations of argue that A. dunniae should include A. Northland, New Zealand (Mollusca: Helicoverpa spp. (Lepidoptera: Noctuidae) tarangaensis, A. pycrofti and A. sp. Gastropoda: Rhytididae: Paryphantinae). on irrigated soybean. Australian Journal of Entomology 44: 293-298. “Motukokako”, whereas you might Molecular Phylogenetics and Evolution, argue that the A. forsythi we are calling 38, 835-842.

Erickson, D.L., Smith, B.D., Clarke, A.C., Huson, D., Kloepper, T., Lockhart, P.J. and Matisoo-Smith, E. (2005). The Rat Path - Sandweiss, D.H. and Tuross, N. (2005). An Steel, M.A. (2005). Reconstruction of Tracing Polynesian migration through rat Asian origin for a 10,000-year-old Reticulate Networks from Gene Trees In DNA. Wild California - The magazine of the domesticated plant in the Americas. Proceedings of the ninth international California Academy of Science. 58(2):16-19. Proceedings of the National Academy of conference in computational molecular Science, USA 102(51): 18315-18320. biology (RECOMB): 233-249. Matisoo-Smith, E., Roberts, K., Welikala, N., Tannock, G., Chester, P., Feek, D. and Gartrell, B. and Hare, K.M. (2005). Mycotic Jeffares, D.C., Mourier, T and Penny, D. Flenley, J. (2005). DNA and pollen from the dermatitis with digital gangrene and (2006). The biology of intron gain and loss. same Lake Core from New Zealand. Pp. 15- osteomyelitis, and protozoal intestinal TRENDS in Genetics 22 (1): 16-22 28 In C.M. Stevenson, J. M. Ramírez Aliaga, parasitism in Marlborough green geckos F.J. Morin, and N. Barbacci (eds) The (Naultinus manukanus). New Zealand Johnson, K.P., Kennedy, M. and McCracken, Reñaca Papers. VI International Conference Veterinary Journal 53(5): 363-367 K.G. (2006). Reinterpreting the Origins of on Easter Island and the Pacific/VI Congreso Flamingo Lice: Cospeciation or Host- internacional sobre Rapa Nui y el Pacífico. Gluckman, P.D., Hanson, M.A., and Switching? Biology Letters 2: 275-278. The Easter Island Foundation, Los Osos. Spencer, H.G. (2005). Predictive adaptive ISBN 1-880636-08-5 responses and human evolution. Trends in Kennedy, M., Holland, B.R., Gray, R.D. and Ecology and Evolution 20: 527-533. Spencer, H.G. (2005). Untangling Long Miller, H.C., Belov, K. and Daugherty, C.H. Branches: Identifying Conflicting (2005). Characterisation of MHC class II Goremykin, V.V., Holland,B., Hirsch-Ernst, Phylogenetic Signals a priori using Spectral genes from an ancient reptile lineage, K.I. and Hellwig, F.H. (2005). Analysis of Analysis, Neighbor-Net, and Consensus Sphenodon (tuatara). Immunogenetics 57: Acorus calamus chloroplast genome and its Networks. Systematic Biology 54:620-633. 883-891. phylogenetic implications. Molecular Biology and Evolution 22: 1813-1822. Kurland, C.G., Collins, L.J., and Penny, D. Morgan-Richards, M. (2005). Chromosome (2006). Genomics and the Irreducible Nature rearrangements are not accompanied by Hare, K.M. and Cree, A. (2005). Natural of Eukaryote Cells. Science 312: 1011-1014. expected genome size change in the tree history of Hoplodactylus stephensi (Reptilia: weta Hemideina thoracica (Orthoptera, Gekkonidae) on Stephens Island, Cook Larson, G., Dobney, K., Albarella, U., Fang, Anostostomatidae). Journal of Orthoptera Strait, New Zealand. New Zealand Journal M., Matisoo-Smith, E., Robins, J., Lowden, Research, 14(2): 143-148. of Ecology 29(1): 137-142. S., Finlayson, H., Brand, T., Willerslev, E., Rowley-Conwy, P., Andersson L. and Ovidiu, P., Lehnebach, C., Johansson, J.T., Hare, K.M., Miller, J.H., Clark, A.G. and Cooper, A. (2005). Worldwide Lockhart, P.J. and Hörandl, E. (2005). Daugherty, C.H. (2005). Muscle lactate phylogeography of wild boar reveals multiple Phylogenetic relationships and biogeography dehydrogenase is not cold-adapted in centres of pig domestication. Science of Ranunculus and allied genera nocturnal lizards from cool-temperate 307:1618-1621. (Ranunculaceae) in the Mediterranean habitats. Comparative Biochemistry and region and in the European Alpine System. Physiology, Part B 142(4): 438-444. Larson, G., Dobney, K., Albarella, U., Taxon 54: 911-930. Matisoo-Smith, E., Robins, J., Lowden, S., Hendy, M.D. (2005). Hadamard Rowley-Conwy, P., Andersson, L. and Penny, D. (2005). An interpretive review of conjugation: an analytic tool for Cooper, A. (2005). Response to the origin of life research. Biology and phylogenetics. Chapter 6, pp 143-177, In Domesticated Pigs in Eastern Indonesia. Philosophy 20:633–671 Mathematics of Evolution and Phylogeny Science 309:381. (O.Gascuel ed), Oxford University Press. Perrie, L.R., Shepherd. L.D. and Brownsey, Lockhart, P.J., Novis, P., Milligan, B.G., P.J. (2005). Asplenium xlucrosum nothosp. Hogg, I.D., Stevens, M.I., Schnabel, K.E. Riden, J., Rambaut, A. and Larkum, A.W.D. Nov.: a sterile hybrid widely and erroneously and Chapman, M.A. (2006). Deeply (2005) Heterotachy and Tree Building: A cultivated as “Aspelium bulbiferum”. Plant divergent lineages of the widespread New Case Study with Plastids and Eubacteria. Syst Evol 250:243-257. Zealand amphipod Paracalliope fluviatilis Mol Biol Evol 23: 40-45. revealed using allozyme and mitochondrial Phillips, M.J. (2006). Sympathy for the Devil. DNA analyses. Freshwater Biology 51: 236- Lockhart, P.J. and Penny, D. (2005). The Nature 440. 248. place of Amborella in the radiation of angiosperms. Trends Plant Sci.10: 201-202. Phillips, M.J., McLenachan, P.A., Down, C., Holland, B. and Schmid, J. (2005). Selecting Gibb, G.C. and Penny, D. (2006). representative model strains. BMC Lockhart, P. and Steel, M. (2005). A tale of Combined nuclear and mitochondrial protein Microbiology 5: 26. two processes. Systematic Biology, 54(6): -coding DNA sequences resolve the 948-951. interrelations of the major Australasian Hörandl, E., Paun, O., Johansson, J.T., marsupial radiations. Systematic Biology 55: Lehnebach, C., Armstrong, T., Chen, L. and McCallum, J., Clarke, A., Pither-Joyce, M., 122-137. Lockhart, P.J. (2005). Phylogenetic Shaw, M., Butler, R., Brash, D., Scheffer, J., relationships and evolutionary traits in Sims, I., van Heusden, S., Shigyo, M. and Robins, J.H., Ross, H.A., Allen, M.S. and Ranunculus s.l. (Ranunculaceae) inferred Havey, M. J. (2006). Genetic mapping of a Matisoo-Smith, E.M. (2006). Sus from ITS sequence analysis Mol Phyl Evol major gene affecting onion bulb fructan bucculentus revisited. Nature 440. 36: 305-327. content. Theoretical and Applied Genetics 112(5): 958-967. Semple, C. and Steel, M. (2006). Unicyclic Huber, K., Moulton, V. and Steel, M. (2005). networks: compatibility and enumeration. Four characters suffice to convexly define a McGaughran, A., Hogg, I.D., Stevens, M.I., IEEE/ACM Transactions on Computational phylogenetic tree. SIAM Journal on Discrete Chadderton, W.L. and Winterbourn, M.J. Biology and Bioinformatics 3(1), 84-91. Mathematics 18(4): 835--843. (2006). Genetic divergence of three freshwater isopod species from southern New Zealand. Journal of Biogeography 33: 23-30.

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