Cyrtandra and other supertramps

Quentin Cronk University of British Columbia Canada

PSI 2009 Acknowledgements

• Jean-Yves Meyer, Pacific Cyrtandra • B.L. “Bill” Burtt, Asian Cyrtandra • Students: Hannah Atkins, Gemma Bramley, Jill Preston • Jim Smith, Boise, Idaho • The organizers, PSI Supertramp Diamond (1974) coined the term ‘supertramp’ for species (birds) of high dispersability that are mainly found on small or isolated — evidence that, in these cases, high dispersability compensates for ecological specialization/ poor competitive ability .

DIAMOND, J. M. 1974. Colonization of exploded volcanic islands by birds: the supertramp strategy. Science 184: 803– 806. Indo-pacific strand flora - supertramps?

• Native - , Ipomoea pes- caprae, Vigna marina, subcordata, taccada, Pisonia grandis • Palaeosynanthropic - tiliaceus, • Neosynanthropic - , Casuarina equisetifolia Types of supertramp

• those in which little speciation has occured (e.g. Thespesia ) • those characterized by at least some radiations ( Bidens , Metrosideros ). Question 1: Patterns of invasion in the Pacific • species widespread among many islands - tectorius (Pandanaceae; Wagner et al., 1990) • multiple radiations on many islands - Psychotria (Rubiaceae; Nepokroeff et al., 2003 • combination of a few widespread species and endemic island species - Scaevola (; Howarth et al., 2003); Metrosideros (Myrtaceae: Percy et al., 2008) • single island lineages - Hillebrandia () Question 2: Number of origins of Pacific by lineages • Single invasion of the Pacific - Metrosideros ( origin in New Zealand); Cyrtandra (origin in SE Asia) • Multiple invasions - Scaevola (Howarth et al., 2003) Cyrtandra

600++ species in Asia and one of the major radiations of the Pacific

• What is the pattern of invasion ? Widespread species? Endemic species?

• How many origins of the Pacific species? One or more? Cyrtandra land Cyrtandra diversity 1

Cyrtandra sp., Borneo Cyrtandra diversity 2

Cyrtandra clarkei (Borneo) Cyrtandra diversity 3

Cyrtandra hawaiiensis x calpidicarpa , photo: J.F. Smith Uniformity of Pacific Cyrtandra

• Foliar sclereids - not found in Pacific species (common in Borneo) • Pollen - Pacific Cyrtandra have uniform pollen type • surface morphology - Pacific Cyrtandra have only the reticulate type (Asian species are heterogeneous - reticulate and striate) • - Pacific species thin walled and white at ripeness (Asian species often thick walled and green or brown at ripeness) Phylogeography of Pacific Cyrtandra

From Cronk et al., 2005 The Pacific lineage

• One colonization of the Pacific • Spread throughout the tropical Polynesian islands • Island specialists, no back-colonization of continental regions, probably poor competitive ability • High dispersability combined with speciation • Conclusion: a supertramp lineage From: Clark et al., 2008 Other islands

Pacific clade Hawaii Cyrtandra in the Pacific

• Most are single island endemics only • High rates of speciation relative to dispersal • Hawaiian Islands - c. 58 endemic species • Marquesas Islands - c. 8 endemic species • Society Islands - c.12 endemic species • Also endemics in , Fiji , Samoa , Tonga , throughout French Dispersal vs speciation

2 Rapid Hundreds - 10 Supertramp species dispersers Thousands - 10 3

4 Moderate 10s of thousands - 10 Lineage dispersers 100s of thousands10 5 supertramps Millions - 10 6 Infrequent Geographically dispersers 10s of millions - 10 7 restricted A new plant phylogeography of the Pacific?

Solve the problems of markers to follow the supertramps:

(a) Conservatism of organellar markers (b) Problems of nuclear markers

The potential rewards are great