Global research on ultramafic (serpentine) ecosystems(8th International Conference on Serpentine Ecologyin Sabah Malaysia) a summary and synthesis

Antony van der EntAEH Nishanta RajakarunaBC Robert BoydD Guillaume Echevarria ERimi RepinF and Dick WilliamsG

ACentre for Mined Land Rehabilitation Sustainable Minerals Institute The University of QueenslandQld Australia

BCollege of the Atlantic 105 Eden Street ME 04609 USACEnvironmental Sciences and Management North-West University Private Bag X6001Potchefstroom 2520 South Africa

DDepartment of Biological Sciences 101 Rouse Life Sciences Bldg Auburn University AL 36849 USAELaboratoire Sols et Environnement UMR 1120 Universiteacute de Lorraine ndash INRA FranceFSabah Parks KK Times Square Coastal Highway 88100 Kota Kinabalu MalaysiaGAustralian Journal of Botany CSIRO Tropical Ecosystems Research Centre AustraliaHCorresponding author Email avanderentuqeduau

Abstract Since 1991 researchers from approximately 45 nations have participated in eight International Conferenceson Serpentine Ecology (ICSE) The Conferences are coordinated by the International Serpentine Ecology Society (ISES)a formal research society whose members study geological pedological biological and applied aspects of ultramafic(serpentine) ecosystems worldwide These conferences have provided an international forum to discuss and synthesisemultidisciplinary research and have provided opportunities for scientists in distinct fields and from different regions of theworld to conduct collaborative and interdisciplinary research The 8th ICSE was hosted by Sabah Parks in Malaysia on theisland of Borneo and attracted the largest delegation to date 174 participants from 31 countries This was the first time anICSE was held in Asia a region that hosts some of the worldrsquos most biodiverse ultramafic ecosystems The presentationsprovided a cross-section of the current status of research in all aspects of ultramafic-biota relations In this Special Issue ofAustralian Journal of Botany (Issues 1ndash2 combined and 3ndash4 combined) we have compiled a selection of papers fromamong the oral and poster presentations to provide insights into recent advances in geoecological and applied studies ofultramafic habitats worldwideHerewe provide a previewof select papers found in this Special Issue and summarise some ofthe contributions made during the 8th ICSE and describe some of the exciting challenges awaiting future research

Received 6 March 2015 accepted 11 March 2015 published online 27 May 2015

Introduction

Ultramafic outcrops (also called lsquoserpentinersquo) are widespreadbut sparse covering roughly 3 of the Earthrsquos surface (Guillotand Hattori 2013) The largest outcrops occur in Cuba NewCaledonia Indonesia the Philippines and Malaysia whereassmaller outcrops are found worldwide mostly along continentalmargins and orogenic belts (Brooks 1987Alexander et al 2007)Soils derived from ultramafic bedrock pose several edaphicchallenges for plant growth including metal toxicity nutrientimbalances and deficiencies and in some cases water stresswith this last feature resulting from the often shallow rocky andexposed nature of the outcrops (Proctor 2003 OrsquoDell andRajakaruna 2011) Ultramafic ecosystems are renowned fortheir high levels of plant diversity and endemism as well as

their unique plantndashhabitat relations (Brooks 1987 Boyd et al2004 2009 Harrison and Rajakaruna 2011) In New Caledonia2150 species occur on ultramafic soils (out of a total of 3371species for the whole flora) of which 83 are restricted to suchsoils (Jaffreacute 1992 Jaffreacute and LrsquoHuillier 2010) whereas in Cuba920 species (approximately one-third of the taxa endemic toCuba) are found exclusively on ultramafic soils (Borhidi1992) Similar restrictions and notable floristic associations arealso found on ultramafic outcrops of theMediterranean region aswell as in Africa Australia and Asia (Brooks 1987 Baker et al1992 Jaffreacute et al 1997 Alexander et al 2007 Chiarucci andBaker 2007 van der Ent et al 2014a) The edaphic challengesassociatedwith these island-like habitats have led to the evolutionof unique ecosystems providing model settings for exploration

CSIRO PUBLISHING

Australian Journal of Botany 2015 63 1ndash16Introduction

httpdxdoiorg101071BT15060

Journal compilation CSIRO 2015 wwwpublishcsiroaujournalsajb

of biological questions at cellular and organismal levels andfor the study of ecosystem-level processes (Harrison andRajakaruna 2011)

International Conferences on Serpentine Ecology(1991ndash2011)

Since 1991 researchers from ~45 nations have participated ineight International Conferences on Serpentine Ecology (ICSE)Conference delegates have come from all corners of the worldincluding Albania Australia Bulgaria Canada China CubaCzech Republic DR Congo France Germany Greece IndiaIran Italy Japan South Korea New Caledonia New ZealandPortugal Russia South Africa Spain Sri Lanka UK andUSA among others The ICSE conferences are coordinated bythe International Serpentine Ecology Society (ISES) a formalresearch society whose members study geological pedologicalbiological and applied aspects of ultramafic ecosystemsworldwide Each conference has highlighted a region withintriguing ultramafic soilndashbiota relations California in 1991(Baker et al 1992) New Caledonia in 1995 (Jaffreacute et al1997) South Africa in 1999 (Balkwill 2001) Cuba in 2003(Boyd et al 2004) Italy in 2006 (Chiarucci and Baker 2007)Maine and eastern Canada in 2008 (Rajakaruna and Boyd2009) and Portugal in 2011 (with a number of articles thatappeared in Plant Ecology and Diversity) These conferenceshave provided an international forum to discuss and synthesisemultidisciplinary research and have provided opportunities forscientists in distinct fields and from different regions of theworld to conduct collaborative and interdisciplinary research

The 8th International Serpentine Ecology Conferencein Sabah Malaysia (2014)

The 8th ICSE was hosted by Sabah Parks in Malaysia on theisland of Borneo The Conference attracted the largest delegationto date 174 participants from 31 countries (Fig 1) TheConference took place over 3 days at the Sutera HarbourResort in Kota Kinabalu followed by 2 days at Kinabalu ParkHeadquarters The Conference was divided into the following sixsessions (1) hyperaccumulator plants and phytotechnologies(2) geology and soils (3) ecology and biogeography (4)physiology and evolution (5) threats and conservation and (6)

diversity systematics and taxonomy In total 37 posterpresentations and 57 oral presentations were delivered duringthe conference covering all aspects of research on ultramaficecosystems The mid-conference fieldtrip was to the abandonedMamut Copper Mine (Fig 2) At the mine delegates had afirst-hand look at the environmental damage caused bydecades of mining a harsh reality worldwide for regionswith metal-rich geologies The importance of such lsquonaturallaboratoriesrsquo for biological ecological evolutionary andapplied research was also emphasised during the excursionImmediately following the conference a 1-day Pacific RimApplication and Grid Middleware Assembly (PRAGMA)special symposium was organised entitled lsquoBiodiversityComputing and Data Infrastructure and Analysisrsquo in which~40 delegates participated After the conference manydelegates took part in post-conference excursions includingclimbs of Mount Kinabalu and Mount Tambuyukon sitesthat are rich with rare and endemic species (van der Ent et al2014b 2015 Aiba et al 2015) During the excursions delegateshad the chance to witness some of the famed iconic speciesfrom Kinabalu Park including the worldrsquos largest flower (theparasitic Rafflesia keithii Rafflesiaceae) the worldrsquos largestcarnivorous pitcher plant (Nepenthes rajah Nepenthaceae) andspectacular orchids such as Paphiopedilum rothschildianum(Orchidaceae) The conference also offered the opportunityfor Sabah Parks to display and disseminate the importantwork it does in conservation and in advancing research onultramafic ecosystems in Kinabalu Park and elsewhere inSabah This was the first time an ISCE was held in Asia aregion that hosts some of the worldrsquos most biodiverseultramafic ecosystems (van der Ent et al 2014a 2015)

Current global research on the ecology and evolutionof ultramafic ecosystems

The 8th International Conference on Serpentine Ecology was aforum for researchers to discuss multidisciplinary research onultramafic ecosystems worldwide Thus the presentationsprovided a cross-section of the current status of research in allaspects of ultramaficndashbiota relations In these Special Issues ofAustralian Journal of Botany (Issues 1ndash2 combined and 3ndash4combined) we have compiled a selection of papers from among

Fig 1 The delegates of the 8th International Conference on Serpentine Ecology in Sabah Malaysia

2 Australian Journal of Botany A van der Ent et al

the oral and poster presentations to provide insights into recentadvances in geoecological and applied studies of ultramafichabitats worldwide Below we provide a preview of selectpapers found in this Special Issue organised by the sessionunder which those papers were presented

Hyperaccumulator plants and phytotechnologies(Session 1)

Ultramafic soils host the greatest number of metalhyperaccumulator plants in the world and these unique plantshave been the subject of floristic physiological evolutionary andapplied research particularly in the past two decades (van der Entet al 2013a Pollard et al 2014) Hyperaccumulators have theremarkable physiological capacity to accumulate heavy metalsandmetalloids in leaf tissues at levels that are orders of magnitudegreater than concentrations found in most other plants (Gall andRajakaruna 2013) It is however a rare phenomenon with only1ndash2 of plant species found on ultramafic soils displaying thisbehaviour A smaller but increasing number of plants are knownas lsquofacultative hyperaccumulatorsrsquo hyperaccumulating heavymetals only when occurring on metal-rich soils but alsooccurring commonly on normal non-metalliferous soils (Pollardet al 2014 McAlister et al 2015) Currently ~450 nickel (Ni)hyperaccumulators are known (van der Ent et al 2013a)Hyperaccumulation is thought to have evolved to interfere withother competing plant species (lsquoelemental allelopathyrsquo) or toprotect against insect herbivores (lsquoelemental herbivorydefencersquo) among other explanations (Boyd 2014) The abilityof hyperaccumulators to extract substantial quantities of Nifrom the soil has sparked the development of phytomining anew technology to extract Ni by harvesting and processing ofhyperaccumulator biomass (Chaney et al 2007 2014) Extensive

ultramafic outcrops in Southeast Asia have the greatest potentialfor future phytomining operations because of the occurrence ofa large number of native Ni-hyperaccumulator species (Reeves2003 Gall and Rajakaruna 2013) and the presence of extensivelateritic Ni-mining operations in the region these have createdpost-mined lands needing rehabilitation that might be usefulphytomining sites (van der Ent et al 2013b)

The session began with keynote presentations by twopioneers in the field of phytoremediation First Baker (2014)reviewed the development of phytotechnologies stressing thatlarge-scale implementation of this green technology is limiteddespite some successful demonstrations in China for arsenic (AsMandal et al 2014) and cadmiumndashzinc (CdndashZn) phytoextraction(Li et al 2014) Second Reeves (2014) focussed on Niphytomining (Keeling et al 2003 Boominathan et al 2004Zhang et al 2014) and what is necessary to see its successfulimplementation He noted areas in need of further research ifphytomining is to reach its potential including improving plantgrowth under metal-rich conditions developing cultivationmethods for agronomic improvement designing managementpractices for dealing with pests and diseases and creating newtechniques to harvest and process biomass to maximise metal-recovery efficiency (Reeves 2014)

Manganese (Mn) hyperaccumulation was highlighted intwo presentations The first was a study on Chengiopanaxsciadophylloides (Araliaceae) from Japan which accumulateda maximum foliar Mn concentration of 23 000mg gndash1 dry weightfrom soils containing low to normal concentrations of Mn(Mizuno et al 2014) The authors obtained a highly purifiedMn compound from solutions made from ashed leaves when thepH was adjusted to 8ndash10 Working with Australian species ofthe genus Gossia (Myrtaceae) Fernando et al (2014) found thatin vivo spatial distributions and subcellular compartmentalisation

Fig 2 Delegates at the Mamut Copper Mine during the mid-conference field trip

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 3

of excessively co-accumulated foliar metals including Mnindicated mediation by specific metal transporters at the cellboundary andor at the tonoplast

Study of relationships of trace element concentrations in soilsand plants and their accumulation kinetics are crucial to designphytoextraction applications that maximise the concentrationof a target element in plant biomass Studies in New Caledoniafound significant effects of leaf age on Ni and Mn accumulationwhich could have implications for producing metal-richbiomass to convert into lsquoecocatalystsrsquo (Losfeld et al 2014a)The concept of lsquoecocatalystsrsquo is based on preparation of novelcatalysts from hyperaccumulator biomass These catalystscan be used in the synthesis of molecules using what hasbeen called lsquogreen chemistryrsquo (Losfeld et al 2014b) in whichthese catalysts replace traditional ones The implementationof lsquoecocatalystsrsquo in the organic chemical industry is a strongencouragement for the development of phytoextractionoperations (Grison and Biton 2014) In New Caledoniaphytoextraction with Geissois pruinosa (Cunoniaceae) a Nihyperaccumulator and two subspecies of Grevillea exul(Proteaceae) both Mn accumulators is underway on minewastes (Losfeld et al 2014c) Ongoing large-scale Niphytomining trials in Albania were reported to yield 100 kgNi handash1 from which a high-value product (Ni ammoniumsulfate hexahydrate) was produced (Zhang et al 2014) Usingtwo native Albanian Ni hyperaccumulators (Alyssum muraleand A markgrafii Brassicaceae) a field cropping regimewas developed that yielded up to 17 t handash1 biomass containing120 kg handash1 Ni (Bani et al 2015) The effects of speciescomposition (mono-cropping or co-cropping) with single andmixtures of Ni hyperaccumulator species (Alyssum muraleNoccaea tymphaea Leptoplax emarginata and Bornmuelleratymphaea all Brassicaceae) were also studied in relation to theefficiency of Ni extraction the results showed that biomass andshoot Ni concentrations (though not significant) of B tymphaeaincreased in the co-cropping system (Rue et al 2015)

Recent discoveries of Ni-hyperaccumulator plants werereported from the Philippines in the islands of PalawanSurigao and Zambales where three species of Phyllanthus(Euphorbiaceae) were found containing gt1 foliar Ni Theyinclude Phyllanthus balgooyi with 7638mg gndash1 Ni andP erythrotrichus and P securinegoides both with gt10 000mggndash1 Ni (Quimado et al 2015) Five new Ni hyperaccumulatorswere also reported from Sri Lanka (Euphorbia heterophylla(Euphorbiaceae) Vernonia cinerea (Asteraceae) Flacourtiaindica (Salicaceae) Olax imbricata (Olacaceae) and Toddaliaasiatica (Rutaceae) Iqbal and Rajakaruna 2014) adding to thethree previously documented for the island by Rajakaruna andBohm (2002) The occurrence of copper (Cu) hyperaccumulationamong some ultramafic plants a phenomenon previouslyreported only for five species from Sri Lanka (Rajakaruna andBaker 2004) was also recently confirmed forMalaysia andBrazil(van der Ent and Reeves 2015) Interestingly a laboratory studybyGhasemi et al (2015a) showed that in the presenceofNiCu isaccumulated in both roots and shoots of Ni-hyperaccumulatingAlyssum spp but not in their non-ultramafic congeners Ghasemiet al (2015a) suggested that Cu hyperaccumulation in thepresence of Ni an abundant metal found in serpentine soilsmay be common In Indonesia foliar elemental concentrations

were studied in Emilia sonchifolia (Asteraceae) grown in top andoverburden soils of the local Ni mines (Tjoa and Barus 2014)Their work showed that although this species produced two- tofive-fold greater shoot biomass when grown in top soilsNi-removal rate was higher in overburden soils (which had ahigher Ni concentration) The chromium (Cr) phytoextractionpotential of Brassica juncea (Brassicaceae) was studied inSri Lanka showing that accumulation was dependent on thegenotype and the ionic concentration of the medium highestfoliar accumulation (3511mg gndash1) occurred when plants weresupplied with 200mgmL1 of Cr(VI) (Wijethunga et al 2014)

The use of plants for rehabilitation of mineral wastes wasdemonstrated with Typha angustifolia (Typhaceae) at the formerMamut Copper Mine (Saibeh et al 2014) and by a study onrelationships among soil microbial properties plant cover andbiomass on ultramafic tailings in SouthAfrica (Smith et al 2014)In the latter case marked differences in soil microbial biomassand community structure were observed between the rockdump and the tailings dam with the highest plant biomassbeing recorded at waste piles of intermediate ages and on themoist eastern aspects of the piles The importance of studyingmicrobial relations in phytoremediation efforts was alsoemphasised in a study examining the potential of Hieraciumpilosella (Asteraceae) for phytoremediation in the presence ofsoil microbes (Ogar et al 2014) This study showed beneficialeffects of microbes on plant growth dry mass of shoots androots increased significantly when plants were inoculated withmycorrhizal fungi and nitrogen-fixing bacteria Seneviratne et al(2015) also showed that the inoculation of ultramafic soilswith bacteria and fungi can enhance soil quality and promoteplant growth in the presence of heavy metals Inoculation withmycorrhizal fungi and addition of P can have plant species-specific growth effects as shown by a study investigatingwhether P addition can stimulate plant growth withoutinhibiting mycorrhizal formation (Amir and Gensous 2014)

Geology and soils (Session 2)

Ultramafic rocks containing the serpentine group of mineralsincluding antigorite chrysotile and lizardite have their originswithin the Earthrsquos upper mantle These rocks often form largemassifs and belts or tabular bodies along continental marginsfaults and shear zones Common ultramafic rock types includeperidotites (including dunite wehrlite harzburgite lherzolite)and the secondary alteration products (serpentinites) formedby their hydration within the Earthrsquos crust via a processknown as serpentinisation (Evans et al 2013) Serpentinisationcreates strongly reducing conditions including fluids enrichedwith hydrogen and methane compounds which chemosyntheticmicrobes can exploit for metabolic energy (Cardace and Hoehler2011 McCollom and Seewald 2013 Cardace et al 2014)Detailed accounts of the origins of serpentinites includingtheir mineralogy petrology weathering and geographicdistribution can be found in Coleman and Jove (1992)Moores (2011) and Hirth and Guillot (2013) Over 60 of theglobal Ni supply comes from Ni laterite ores produced from theintensiveweathering of serpentinites found under humid tropicalconditions (Butt andCluzel 2013)Ultramafic soils areweatheredproducts of serpentinite and other ultramafic rocks Ultramafic

4 Australian Journal of Botany A van der Ent et al

bedrock type drainage conditions and weathering intensity allcontribute to differences in pedogenesis and availability ofelements including Ni (Echevarria 2014) For exampleultramafic soils formed by the weathering of peridotite orserpentinite (ultramafic rocks that are chemically similar yetmineralogically distinct) show appreciable differences ingeomorphic and pedologic features (Alexander and DuShey2011) For a summary of the developmental processes ofultramafic soils see Brooks (1987) and Alexander et al(2007) Ultramafic soils are generally deficient in plantessential nutrients such as nitrogen (N) phosphorus (P)potassium (K) and sulfur (S) have a calcium magnesium(Ca Mg) ratio lt1 and have elevated concentrations of heavymetals such as Ni cobalt (Co) Cd and Cr Although physicalfeatures of ultramafic soils can vary considerably from site to site(and within a site) ultramafic soils are often found in open steeplandscapes with substrates that are generally shallow and rockyand that often have reduced capacity for moisture retentionExtensive mining in regions overlying ultramafic bedrockhas led to degraded landscapes needing restoration Carefulmanagement of the topsoil (extracted before the miningdisturbance) can maximise biodiversity on newly restored soilsbecause the topsoil can reintroduce both native seeds andmicrobes to a site under restoration (Bordez et al 2014) Astudy by Mori (2014) examined the potential for greenhousegas emissions (CO2 CH4 N2O) from undisturbed forest soilson different parent materials (sedimentary rock and serpentinerock) along an altitudinal gradient (700 1700 2700 and 3100masl) on Mount Kinabalu Borneo His work suggested thatcarbon fluxes decrease with increasing altitude implying thatlower temperature inhibits microbial activities N2O emissionshowever were not altitude-dependent but were lower onultramafic soils than on sedimentary soils likely because ofthe lower N availability and closed N cycle in ultramafic soilsincluding lower inorganic N nitrification potential microbial Nand higher NH4

+ NO3ndash ratio Whether ultramafic rocks can save

the world from increased carbon emissions has also been a focusof study (Power et al 2013) Mg-rich serpentinite and relatedrocks can react withwater andCO2 to formmagnesium carbonateplus silica thus sequestering potentially damaging CO2

emissions (Maroto-Valer et al 2005 Yang et al 2008) Thistechnique if implemented might have a drastic impact on theunique biota of ultramafic areas

Ecology and biogeography (Session 3)

The unique features of ultramafic habitats can have importanteffects on organismal interactions providing many stimulatingavenues for researchUltramafic sitesmay contain unique speciesof bacteria fungi plants and animals and these often areassembled into communities distinct from surrounding areasFloras of ultramafic sites have attracted considerable attentionat past International Conferences on Serpentine Ecology and the2014Conferencewasnoexception In this session comparisonofultramafic and non-ultramafic forests on Mount Kinabalu (Aibaet al 2015) description of the ultramafic flora of Sri Lanka (Iqbaland Rajakaruna 2014) and a review (Siebert 2014) of SouthAfrican ultramafic-ecology research (which showed that muchwork there has focussed on species descriptions and investigation

of diversity patterns) added to our global knowledge of ultramaficplant ecology Several contributions also added to our floristicknowledge for ultramafic sites in Malaysia (Damit et al 2014Majapun et al 2014 Pereira et al 2014 Sabran et al 2014)including studies conducted at the venue for the poster sessionKinabalu Park (Karim and van der Ent 2014 Sumail and van derEnt 2014) Other presentations contributed floristic informationfrom South Africa (Frisby et al 2014) Japan (Mizuno andKirihata 2015) and Iran (Mohtadi et al 2014) Burgess et al(2015a) documented successional changes that have occurred onultramafic grasslands in the Mid-Atlantic region of the USAshowing that grass-dominated sites have shifted to forests over adecades-long time frame

Rajakaruna and Boyd (2014) pointed out that ultramaficfaunas are little studied compared with ultramafic florasContributions from past International Conferences onSerpentine Ecology (Chazeau 1997 Jourdan 1997 Cantildeameroet al 2004) have added to our knowledge of fauna This traditioncontinued at the 2014 Conference with presentations by Chunget al (2014) on insects from ultramafic sites in a Malaysianforest reserve and Homathevi et al (2014) on termite diversityon ultramafic sites on Mount Tambuyukon Sabah Lichens ofserpentine areas are receiving more study in part because ofwork stimulated bypresentations at past conferences (eg Paukov2009) The 2014 Conference received new contributions on thelichen floras of ultramafic areas in Europe (Paukov et al 2014Favero-Longo et al 2015) and the north-eastern US (Medeiroset al 2014) In contrast to plants and lichens bacteria andmycorrhizal communities seem less affected by ultramaficsoils For example Oline (2006) found little evidence thatthere is a unique ultramafic-soil bacterial flora Venter et al(2015) examined soil algae and cyanoprokaryotes in ultramaficand non-ultramafic soils in South Africa and reached a similarconclusion but did discover that some species were unique to theultramafic communities In a review produced from the 2011conference Southworth et al (2014) found little evidence thatmycorrhizal fungal communities are edaphically specialisedThis may stem from ectomycorrhizal fungi being less inhibitedby edaphic stressors than are plants growing in ultramafic soils(Branco and Ree 2010)

One fascinating component of many ultramafic floras aremetal-hyperaccumulator plants Researchers at the 2014conference continued to catalogue these species describe theirassociates and explore their physiology and ecology van der Entet al (2015) added 19 new Ni-hyperaccumulator species tothe five previously recorded from ultramafic soils in Sabah(Malaysia) illustrating the potential for new discoveries of theseunusual plants Mesjasz-Przybylowicz and Przybylowicz (2014)highlighted intensive and wide-ranging research accomplishedin South Africa on several Ni-hyperaccumulator plant speciesincluding investigations of the mycorrhizal and herbivoreassociates of these plants (also see Mesjasz-Przybyłowicz andPrzybyłowicz 2011) Metal-hyperaccumulator plants may usemetals for defence or the metals may have other functions(such as elemental allelopathy or drought resistance) as recentlyreviewed by Boyd (2014) In a contribution from the 2006conference Boyd (2007) pointed out that most studies of metalhyperaccumulation as a plant defence have focussed on whethermetals increase plant resistance to herbivore attack But tolerance

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 5

of herbivory (ability to withstand damage) may be anotherplant defence tactic affected by metal hyperaccumulationPalomino et al (2007) first addressed this question in a paperpublished in the 2006 conference proceedings findingsignificantly increased tolerance by Noccaea fendleri subspglauca to herbivore damage when plants hyperaccumulated ZnMincey and Boyd (2014) showed elevated tolerance of herbivoryby a Ni-hyperaccumulator species (Streptanthus polygaloidesBrassicaceae) confirming and extending the earlier work with Zn

Finally the patchiness of ultramafic habitats at many sitescan have important impacts on ecological relationships (egpollination seed dispersal) that can affect gene flow andspeciation For example research presented at the 2011conference (Spasojevic et al 2014) explored patterns of seed-dispersal syndromes in Californian ultramafic and non-ultramaficfloras including the influence of ultramafic patch size Substratetype significantly influenced seed dispersal by vertebrates andwind but patch size did not have an important effect At the2014 Conference the patchiness of ultramafic habitats wasevident by presentations from many locations including forexample South Africa Malaysia and the Philippines In thissession Porembski (2014) pointed out similar patchiness ofother geological substrates such as granitoid and ironstoneoutcrops that also results in unique communities Morerecently other edaphic island communities such as those foundon gabbro (Medeiros et al 2015) and gypsum (Moore et al 2014Escudero et al 2015) have also received close attention as settingscomparable to ultramafic habitats for examining ecological andevolutionary theory (Harrison and Rajakaruna 2011) Some of thefloristic works discussed above have important biogeographicaspects because they document the distributions of speciesacross these patchy edaphic landscapes Other contributions tothis session targeted specific biogeographic relationshipsOne such biogeographic concept Rapoportrsquos rule (Veter et al2013) was the topic of the presentation by Whitman and Russo(2014) Rapoportrsquos rule states that the size of a speciesrsquo rangeincreases as altitude or latitude increases andWhitman and Russo(2014) found that the rule applied to soil-generalist species onMount Kinabalu but not to ultramafic-soil specialists Theysuggested that edaphic specialisation limits a species range ofclimatic tolerance (also see Fernandez-Going 2014) A study byGhasemi et al (2015b) examined environmental-stress sensitivityunder different Ca Mg ratios by a ultramafic endemic and itsnon-ultramafic congener Their study suggested that underhigher Ca Mg ratios (characteristic of non-ultramafic soil) theultramafic endemic has reduced tolerance to higher N andincreased temperature The implication of this study is thatultramafic endemics may be more susceptible to human-drivenclimate change-associated stressors such as atmospheric Ndeposition (Pasari et al 2014) than non-ultramafic species Thepresentation by Burge (2014) also addressed the impact ofultramafic soils on plant climatic niches Focusing on soilgeneralists from California USA this study examined whetherultramafic soils allow plants to move outside their climatic nichesThis was indeed the case for both upper and lower limits of speciesdistributions It was suggested that at low elevations ultramaficsoilsprovidea refuge fromcompetitionwhereasathighelevationsthe effect of cold is magnified by infertile ultramafic soils (Burgeand Salk 2014)

Physiology and evolution (Session 4)

Because of the high degree of abiotic stress on ultramafic sitesultramafic-associated biota are model organisms for the study ofadaptation (OrsquoDell and Rajakaruna 2011) The low Ca Mg ratioof ultramafic soils is a major challenge to plant growth andmechanisms to deal with this nutrient imbalance are key toultramafic tolerance These mechanisms include tolerance ofhigh concentrations of soil Mg reduced absorption of Mg orhigher absorption of Ca (Palm and Van Volkenburgh 2014) Inaddition low macronutrient concentrations high concentrationsof some heavy metals and low water availability are alsoimportant factors that must be tolerated by ultramafic plantsSeveral poster and oral presentations at the conference addressedultramafic tolerance with respect to heavy metals and othernutrient imbalances in plants (Doronila et al 2014 Echevarriaet al 2014 Pollard and Smith 2014 Teptina and Paukov 2014Ghaderian et al 2015 Hendry et al 2015) and lichens (Paukovet al 2014 Favero-Longo et al 2015) Although studiesgenerally focus on the effects of a single stress factor thesestress factors are likely to have combined effects (Von Wettberget al 2014) as in the case for Ca Mg ratio and water availabilityon traits of Mimulus guttatus (Phrymaceae Murren et al 2006Selby et al 2014) Presentations also focussed on elementaluptake and localisation in tissues of plants tolerant ofultramafic and other chemically imbalanced soils (Kosugiet al 2015 Mizuno and Kirihata 2015 Pavlova et al 2015)including the use of advanced techniques such as micro-PIXEin mapping regions of elemental concentration within planttissue (Przybyłowicz et al 2005 2014) Tolerance of highconcentrations of heavy metals including molecularmechanisms underlying metal tolerance has also receivedattention (Janssens et al 2009 Gall and Rajakaruna 2013Viehweger 2014) There has been considerable effort tounderstand the linkage between ultramafic-soil stress factorsfor plants (such as metals nutrients water) and the genes thatunderlie adaptations to them (Von Wettberg et al 2014) Recentadvances in molecular biology and genomics have providedpowerful tools to examine the genetic bases for adaptation toultramafic soils (Von Wettberg and Wright 2011 Selby et al2014) In Arabidopsis for example polymorphisms for traitsinvolved in Ca Mg tolerance and heavy-metal detoxificationhave been detected in ultramafic and non-ultramafic populations(Turner et al 2008 2010) suggesting that parallel ecologicaladaptations (Rajakaruna et al 2003 Ostevik et al 2012) canoccur via the differentiation of the same polymorphism atultramafic-tolerant loci in geographically distinct ultramafic-tolerant populations An often used approach for identifyinggenes involved in ultramafic adaptation is the study ofquantitative trait loci (QTL Bratteler et al 2006 Murren et al2006 Wu et al 2008) Recent advances in various fields ofmolecular biology may allow for even newer approaches tostudying ultramafic-plant physiology and the underlyinggenetics of adaptations (Von Wettberg and Wright 2011Selby et al 2014)

Ultramafic plants and their associated biota are also idealmodel systems to explore the role of divergent natural selectionin speciation (Rajakaruna 2004 Kay et al 2011) Intraspecificvariation is central to divergence Population-level variation for

6 Australian Journal of Botany A van der Ent et al

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

the oral and poster presentations to provide insights into recentadvances in geoecological and applied studies of ultramafichabitats worldwide Below we provide a preview of selectpapers found in this Special Issue organised by the sessionunder which those papers were presented

Hyperaccumulator plants and phytotechnologies(Session 1)

Ultramafic soils host the greatest number of metalhyperaccumulator plants in the world and these unique plantshave been the subject of floristic physiological evolutionary andapplied research particularly in the past two decades (van der Entet al 2013a Pollard et al 2014) Hyperaccumulators have theremarkable physiological capacity to accumulate heavy metalsandmetalloids in leaf tissues at levels that are orders of magnitudegreater than concentrations found in most other plants (Gall andRajakaruna 2013) It is however a rare phenomenon with only1ndash2 of plant species found on ultramafic soils displaying thisbehaviour A smaller but increasing number of plants are knownas lsquofacultative hyperaccumulatorsrsquo hyperaccumulating heavymetals only when occurring on metal-rich soils but alsooccurring commonly on normal non-metalliferous soils (Pollardet al 2014 McAlister et al 2015) Currently ~450 nickel (Ni)hyperaccumulators are known (van der Ent et al 2013a)Hyperaccumulation is thought to have evolved to interfere withother competing plant species (lsquoelemental allelopathyrsquo) or toprotect against insect herbivores (lsquoelemental herbivorydefencersquo) among other explanations (Boyd 2014) The abilityof hyperaccumulators to extract substantial quantities of Nifrom the soil has sparked the development of phytomining anew technology to extract Ni by harvesting and processing ofhyperaccumulator biomass (Chaney et al 2007 2014) Extensive

ultramafic outcrops in Southeast Asia have the greatest potentialfor future phytomining operations because of the occurrence ofa large number of native Ni-hyperaccumulator species (Reeves2003 Gall and Rajakaruna 2013) and the presence of extensivelateritic Ni-mining operations in the region these have createdpost-mined lands needing rehabilitation that might be usefulphytomining sites (van der Ent et al 2013b)

The session began with keynote presentations by twopioneers in the field of phytoremediation First Baker (2014)reviewed the development of phytotechnologies stressing thatlarge-scale implementation of this green technology is limiteddespite some successful demonstrations in China for arsenic (AsMandal et al 2014) and cadmiumndashzinc (CdndashZn) phytoextraction(Li et al 2014) Second Reeves (2014) focussed on Niphytomining (Keeling et al 2003 Boominathan et al 2004Zhang et al 2014) and what is necessary to see its successfulimplementation He noted areas in need of further research ifphytomining is to reach its potential including improving plantgrowth under metal-rich conditions developing cultivationmethods for agronomic improvement designing managementpractices for dealing with pests and diseases and creating newtechniques to harvest and process biomass to maximise metal-recovery efficiency (Reeves 2014)

Manganese (Mn) hyperaccumulation was highlighted intwo presentations The first was a study on Chengiopanaxsciadophylloides (Araliaceae) from Japan which accumulateda maximum foliar Mn concentration of 23 000mg gndash1 dry weightfrom soils containing low to normal concentrations of Mn(Mizuno et al 2014) The authors obtained a highly purifiedMn compound from solutions made from ashed leaves when thepH was adjusted to 8ndash10 Working with Australian species ofthe genus Gossia (Myrtaceae) Fernando et al (2014) found thatin vivo spatial distributions and subcellular compartmentalisation

Fig 2 Delegates at the Mamut Copper Mine during the mid-conference field trip

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 3

of excessively co-accumulated foliar metals including Mnindicated mediation by specific metal transporters at the cellboundary andor at the tonoplast

Study of relationships of trace element concentrations in soilsand plants and their accumulation kinetics are crucial to designphytoextraction applications that maximise the concentrationof a target element in plant biomass Studies in New Caledoniafound significant effects of leaf age on Ni and Mn accumulationwhich could have implications for producing metal-richbiomass to convert into lsquoecocatalystsrsquo (Losfeld et al 2014a)The concept of lsquoecocatalystsrsquo is based on preparation of novelcatalysts from hyperaccumulator biomass These catalystscan be used in the synthesis of molecules using what hasbeen called lsquogreen chemistryrsquo (Losfeld et al 2014b) in whichthese catalysts replace traditional ones The implementationof lsquoecocatalystsrsquo in the organic chemical industry is a strongencouragement for the development of phytoextractionoperations (Grison and Biton 2014) In New Caledoniaphytoextraction with Geissois pruinosa (Cunoniaceae) a Nihyperaccumulator and two subspecies of Grevillea exul(Proteaceae) both Mn accumulators is underway on minewastes (Losfeld et al 2014c) Ongoing large-scale Niphytomining trials in Albania were reported to yield 100 kgNi handash1 from which a high-value product (Ni ammoniumsulfate hexahydrate) was produced (Zhang et al 2014) Usingtwo native Albanian Ni hyperaccumulators (Alyssum muraleand A markgrafii Brassicaceae) a field cropping regimewas developed that yielded up to 17 t handash1 biomass containing120 kg handash1 Ni (Bani et al 2015) The effects of speciescomposition (mono-cropping or co-cropping) with single andmixtures of Ni hyperaccumulator species (Alyssum muraleNoccaea tymphaea Leptoplax emarginata and Bornmuelleratymphaea all Brassicaceae) were also studied in relation to theefficiency of Ni extraction the results showed that biomass andshoot Ni concentrations (though not significant) of B tymphaeaincreased in the co-cropping system (Rue et al 2015)

Recent discoveries of Ni-hyperaccumulator plants werereported from the Philippines in the islands of PalawanSurigao and Zambales where three species of Phyllanthus(Euphorbiaceae) were found containing gt1 foliar Ni Theyinclude Phyllanthus balgooyi with 7638mg gndash1 Ni andP erythrotrichus and P securinegoides both with gt10 000mggndash1 Ni (Quimado et al 2015) Five new Ni hyperaccumulatorswere also reported from Sri Lanka (Euphorbia heterophylla(Euphorbiaceae) Vernonia cinerea (Asteraceae) Flacourtiaindica (Salicaceae) Olax imbricata (Olacaceae) and Toddaliaasiatica (Rutaceae) Iqbal and Rajakaruna 2014) adding to thethree previously documented for the island by Rajakaruna andBohm (2002) The occurrence of copper (Cu) hyperaccumulationamong some ultramafic plants a phenomenon previouslyreported only for five species from Sri Lanka (Rajakaruna andBaker 2004) was also recently confirmed forMalaysia andBrazil(van der Ent and Reeves 2015) Interestingly a laboratory studybyGhasemi et al (2015a) showed that in the presenceofNiCu isaccumulated in both roots and shoots of Ni-hyperaccumulatingAlyssum spp but not in their non-ultramafic congeners Ghasemiet al (2015a) suggested that Cu hyperaccumulation in thepresence of Ni an abundant metal found in serpentine soilsmay be common In Indonesia foliar elemental concentrations

were studied in Emilia sonchifolia (Asteraceae) grown in top andoverburden soils of the local Ni mines (Tjoa and Barus 2014)Their work showed that although this species produced two- tofive-fold greater shoot biomass when grown in top soilsNi-removal rate was higher in overburden soils (which had ahigher Ni concentration) The chromium (Cr) phytoextractionpotential of Brassica juncea (Brassicaceae) was studied inSri Lanka showing that accumulation was dependent on thegenotype and the ionic concentration of the medium highestfoliar accumulation (3511mg gndash1) occurred when plants weresupplied with 200mgmL1 of Cr(VI) (Wijethunga et al 2014)

The use of plants for rehabilitation of mineral wastes wasdemonstrated with Typha angustifolia (Typhaceae) at the formerMamut Copper Mine (Saibeh et al 2014) and by a study onrelationships among soil microbial properties plant cover andbiomass on ultramafic tailings in SouthAfrica (Smith et al 2014)In the latter case marked differences in soil microbial biomassand community structure were observed between the rockdump and the tailings dam with the highest plant biomassbeing recorded at waste piles of intermediate ages and on themoist eastern aspects of the piles The importance of studyingmicrobial relations in phytoremediation efforts was alsoemphasised in a study examining the potential of Hieraciumpilosella (Asteraceae) for phytoremediation in the presence ofsoil microbes (Ogar et al 2014) This study showed beneficialeffects of microbes on plant growth dry mass of shoots androots increased significantly when plants were inoculated withmycorrhizal fungi and nitrogen-fixing bacteria Seneviratne et al(2015) also showed that the inoculation of ultramafic soilswith bacteria and fungi can enhance soil quality and promoteplant growth in the presence of heavy metals Inoculation withmycorrhizal fungi and addition of P can have plant species-specific growth effects as shown by a study investigatingwhether P addition can stimulate plant growth withoutinhibiting mycorrhizal formation (Amir and Gensous 2014)

Geology and soils (Session 2)

Ultramafic rocks containing the serpentine group of mineralsincluding antigorite chrysotile and lizardite have their originswithin the Earthrsquos upper mantle These rocks often form largemassifs and belts or tabular bodies along continental marginsfaults and shear zones Common ultramafic rock types includeperidotites (including dunite wehrlite harzburgite lherzolite)and the secondary alteration products (serpentinites) formedby their hydration within the Earthrsquos crust via a processknown as serpentinisation (Evans et al 2013) Serpentinisationcreates strongly reducing conditions including fluids enrichedwith hydrogen and methane compounds which chemosyntheticmicrobes can exploit for metabolic energy (Cardace and Hoehler2011 McCollom and Seewald 2013 Cardace et al 2014)Detailed accounts of the origins of serpentinites includingtheir mineralogy petrology weathering and geographicdistribution can be found in Coleman and Jove (1992)Moores (2011) and Hirth and Guillot (2013) Over 60 of theglobal Ni supply comes from Ni laterite ores produced from theintensiveweathering of serpentinites found under humid tropicalconditions (Butt andCluzel 2013)Ultramafic soils areweatheredproducts of serpentinite and other ultramafic rocks Ultramafic

4 Australian Journal of Botany A van der Ent et al

bedrock type drainage conditions and weathering intensity allcontribute to differences in pedogenesis and availability ofelements including Ni (Echevarria 2014) For exampleultramafic soils formed by the weathering of peridotite orserpentinite (ultramafic rocks that are chemically similar yetmineralogically distinct) show appreciable differences ingeomorphic and pedologic features (Alexander and DuShey2011) For a summary of the developmental processes ofultramafic soils see Brooks (1987) and Alexander et al(2007) Ultramafic soils are generally deficient in plantessential nutrients such as nitrogen (N) phosphorus (P)potassium (K) and sulfur (S) have a calcium magnesium(Ca Mg) ratio lt1 and have elevated concentrations of heavymetals such as Ni cobalt (Co) Cd and Cr Although physicalfeatures of ultramafic soils can vary considerably from site to site(and within a site) ultramafic soils are often found in open steeplandscapes with substrates that are generally shallow and rockyand that often have reduced capacity for moisture retentionExtensive mining in regions overlying ultramafic bedrockhas led to degraded landscapes needing restoration Carefulmanagement of the topsoil (extracted before the miningdisturbance) can maximise biodiversity on newly restored soilsbecause the topsoil can reintroduce both native seeds andmicrobes to a site under restoration (Bordez et al 2014) Astudy by Mori (2014) examined the potential for greenhousegas emissions (CO2 CH4 N2O) from undisturbed forest soilson different parent materials (sedimentary rock and serpentinerock) along an altitudinal gradient (700 1700 2700 and 3100masl) on Mount Kinabalu Borneo His work suggested thatcarbon fluxes decrease with increasing altitude implying thatlower temperature inhibits microbial activities N2O emissionshowever were not altitude-dependent but were lower onultramafic soils than on sedimentary soils likely because ofthe lower N availability and closed N cycle in ultramafic soilsincluding lower inorganic N nitrification potential microbial Nand higher NH4

+ NO3ndash ratio Whether ultramafic rocks can save

the world from increased carbon emissions has also been a focusof study (Power et al 2013) Mg-rich serpentinite and relatedrocks can react withwater andCO2 to formmagnesium carbonateplus silica thus sequestering potentially damaging CO2

emissions (Maroto-Valer et al 2005 Yang et al 2008) Thistechnique if implemented might have a drastic impact on theunique biota of ultramafic areas

Ecology and biogeography (Session 3)

The unique features of ultramafic habitats can have importanteffects on organismal interactions providing many stimulatingavenues for researchUltramafic sitesmay contain unique speciesof bacteria fungi plants and animals and these often areassembled into communities distinct from surrounding areasFloras of ultramafic sites have attracted considerable attentionat past International Conferences on Serpentine Ecology and the2014Conferencewasnoexception In this session comparisonofultramafic and non-ultramafic forests on Mount Kinabalu (Aibaet al 2015) description of the ultramafic flora of Sri Lanka (Iqbaland Rajakaruna 2014) and a review (Siebert 2014) of SouthAfrican ultramafic-ecology research (which showed that muchwork there has focussed on species descriptions and investigation

of diversity patterns) added to our global knowledge of ultramaficplant ecology Several contributions also added to our floristicknowledge for ultramafic sites in Malaysia (Damit et al 2014Majapun et al 2014 Pereira et al 2014 Sabran et al 2014)including studies conducted at the venue for the poster sessionKinabalu Park (Karim and van der Ent 2014 Sumail and van derEnt 2014) Other presentations contributed floristic informationfrom South Africa (Frisby et al 2014) Japan (Mizuno andKirihata 2015) and Iran (Mohtadi et al 2014) Burgess et al(2015a) documented successional changes that have occurred onultramafic grasslands in the Mid-Atlantic region of the USAshowing that grass-dominated sites have shifted to forests over adecades-long time frame

Rajakaruna and Boyd (2014) pointed out that ultramaficfaunas are little studied compared with ultramafic florasContributions from past International Conferences onSerpentine Ecology (Chazeau 1997 Jourdan 1997 Cantildeameroet al 2004) have added to our knowledge of fauna This traditioncontinued at the 2014 Conference with presentations by Chunget al (2014) on insects from ultramafic sites in a Malaysianforest reserve and Homathevi et al (2014) on termite diversityon ultramafic sites on Mount Tambuyukon Sabah Lichens ofserpentine areas are receiving more study in part because ofwork stimulated bypresentations at past conferences (eg Paukov2009) The 2014 Conference received new contributions on thelichen floras of ultramafic areas in Europe (Paukov et al 2014Favero-Longo et al 2015) and the north-eastern US (Medeiroset al 2014) In contrast to plants and lichens bacteria andmycorrhizal communities seem less affected by ultramaficsoils For example Oline (2006) found little evidence thatthere is a unique ultramafic-soil bacterial flora Venter et al(2015) examined soil algae and cyanoprokaryotes in ultramaficand non-ultramafic soils in South Africa and reached a similarconclusion but did discover that some species were unique to theultramafic communities In a review produced from the 2011conference Southworth et al (2014) found little evidence thatmycorrhizal fungal communities are edaphically specialisedThis may stem from ectomycorrhizal fungi being less inhibitedby edaphic stressors than are plants growing in ultramafic soils(Branco and Ree 2010)

One fascinating component of many ultramafic floras aremetal-hyperaccumulator plants Researchers at the 2014conference continued to catalogue these species describe theirassociates and explore their physiology and ecology van der Entet al (2015) added 19 new Ni-hyperaccumulator species tothe five previously recorded from ultramafic soils in Sabah(Malaysia) illustrating the potential for new discoveries of theseunusual plants Mesjasz-Przybylowicz and Przybylowicz (2014)highlighted intensive and wide-ranging research accomplishedin South Africa on several Ni-hyperaccumulator plant speciesincluding investigations of the mycorrhizal and herbivoreassociates of these plants (also see Mesjasz-Przybyłowicz andPrzybyłowicz 2011) Metal-hyperaccumulator plants may usemetals for defence or the metals may have other functions(such as elemental allelopathy or drought resistance) as recentlyreviewed by Boyd (2014) In a contribution from the 2006conference Boyd (2007) pointed out that most studies of metalhyperaccumulation as a plant defence have focussed on whethermetals increase plant resistance to herbivore attack But tolerance

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 5

of herbivory (ability to withstand damage) may be anotherplant defence tactic affected by metal hyperaccumulationPalomino et al (2007) first addressed this question in a paperpublished in the 2006 conference proceedings findingsignificantly increased tolerance by Noccaea fendleri subspglauca to herbivore damage when plants hyperaccumulated ZnMincey and Boyd (2014) showed elevated tolerance of herbivoryby a Ni-hyperaccumulator species (Streptanthus polygaloidesBrassicaceae) confirming and extending the earlier work with Zn

Finally the patchiness of ultramafic habitats at many sitescan have important impacts on ecological relationships (egpollination seed dispersal) that can affect gene flow andspeciation For example research presented at the 2011conference (Spasojevic et al 2014) explored patterns of seed-dispersal syndromes in Californian ultramafic and non-ultramaficfloras including the influence of ultramafic patch size Substratetype significantly influenced seed dispersal by vertebrates andwind but patch size did not have an important effect At the2014 Conference the patchiness of ultramafic habitats wasevident by presentations from many locations including forexample South Africa Malaysia and the Philippines In thissession Porembski (2014) pointed out similar patchiness ofother geological substrates such as granitoid and ironstoneoutcrops that also results in unique communities Morerecently other edaphic island communities such as those foundon gabbro (Medeiros et al 2015) and gypsum (Moore et al 2014Escudero et al 2015) have also received close attention as settingscomparable to ultramafic habitats for examining ecological andevolutionary theory (Harrison and Rajakaruna 2011) Some of thefloristic works discussed above have important biogeographicaspects because they document the distributions of speciesacross these patchy edaphic landscapes Other contributions tothis session targeted specific biogeographic relationshipsOne such biogeographic concept Rapoportrsquos rule (Veter et al2013) was the topic of the presentation by Whitman and Russo(2014) Rapoportrsquos rule states that the size of a speciesrsquo rangeincreases as altitude or latitude increases andWhitman and Russo(2014) found that the rule applied to soil-generalist species onMount Kinabalu but not to ultramafic-soil specialists Theysuggested that edaphic specialisation limits a species range ofclimatic tolerance (also see Fernandez-Going 2014) A study byGhasemi et al (2015b) examined environmental-stress sensitivityunder different Ca Mg ratios by a ultramafic endemic and itsnon-ultramafic congener Their study suggested that underhigher Ca Mg ratios (characteristic of non-ultramafic soil) theultramafic endemic has reduced tolerance to higher N andincreased temperature The implication of this study is thatultramafic endemics may be more susceptible to human-drivenclimate change-associated stressors such as atmospheric Ndeposition (Pasari et al 2014) than non-ultramafic species Thepresentation by Burge (2014) also addressed the impact ofultramafic soils on plant climatic niches Focusing on soilgeneralists from California USA this study examined whetherultramafic soils allow plants to move outside their climatic nichesThis was indeed the case for both upper and lower limits of speciesdistributions It was suggested that at low elevations ultramaficsoilsprovidea refuge fromcompetitionwhereasathighelevationsthe effect of cold is magnified by infertile ultramafic soils (Burgeand Salk 2014)

Physiology and evolution (Session 4)

Because of the high degree of abiotic stress on ultramafic sitesultramafic-associated biota are model organisms for the study ofadaptation (OrsquoDell and Rajakaruna 2011) The low Ca Mg ratioof ultramafic soils is a major challenge to plant growth andmechanisms to deal with this nutrient imbalance are key toultramafic tolerance These mechanisms include tolerance ofhigh concentrations of soil Mg reduced absorption of Mg orhigher absorption of Ca (Palm and Van Volkenburgh 2014) Inaddition low macronutrient concentrations high concentrationsof some heavy metals and low water availability are alsoimportant factors that must be tolerated by ultramafic plantsSeveral poster and oral presentations at the conference addressedultramafic tolerance with respect to heavy metals and othernutrient imbalances in plants (Doronila et al 2014 Echevarriaet al 2014 Pollard and Smith 2014 Teptina and Paukov 2014Ghaderian et al 2015 Hendry et al 2015) and lichens (Paukovet al 2014 Favero-Longo et al 2015) Although studiesgenerally focus on the effects of a single stress factor thesestress factors are likely to have combined effects (Von Wettberget al 2014) as in the case for Ca Mg ratio and water availabilityon traits of Mimulus guttatus (Phrymaceae Murren et al 2006Selby et al 2014) Presentations also focussed on elementaluptake and localisation in tissues of plants tolerant ofultramafic and other chemically imbalanced soils (Kosugiet al 2015 Mizuno and Kirihata 2015 Pavlova et al 2015)including the use of advanced techniques such as micro-PIXEin mapping regions of elemental concentration within planttissue (Przybyłowicz et al 2005 2014) Tolerance of highconcentrations of heavy metals including molecularmechanisms underlying metal tolerance has also receivedattention (Janssens et al 2009 Gall and Rajakaruna 2013Viehweger 2014) There has been considerable effort tounderstand the linkage between ultramafic-soil stress factorsfor plants (such as metals nutrients water) and the genes thatunderlie adaptations to them (Von Wettberg et al 2014) Recentadvances in molecular biology and genomics have providedpowerful tools to examine the genetic bases for adaptation toultramafic soils (Von Wettberg and Wright 2011 Selby et al2014) In Arabidopsis for example polymorphisms for traitsinvolved in Ca Mg tolerance and heavy-metal detoxificationhave been detected in ultramafic and non-ultramafic populations(Turner et al 2008 2010) suggesting that parallel ecologicaladaptations (Rajakaruna et al 2003 Ostevik et al 2012) canoccur via the differentiation of the same polymorphism atultramafic-tolerant loci in geographically distinct ultramafic-tolerant populations An often used approach for identifyinggenes involved in ultramafic adaptation is the study ofquantitative trait loci (QTL Bratteler et al 2006 Murren et al2006 Wu et al 2008) Recent advances in various fields ofmolecular biology may allow for even newer approaches tostudying ultramafic-plant physiology and the underlyinggenetics of adaptations (Von Wettberg and Wright 2011Selby et al 2014)

Ultramafic plants and their associated biota are also idealmodel systems to explore the role of divergent natural selectionin speciation (Rajakaruna 2004 Kay et al 2011) Intraspecificvariation is central to divergence Population-level variation for

6 Australian Journal of Botany A van der Ent et al

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

of excessively co-accumulated foliar metals including Mnindicated mediation by specific metal transporters at the cellboundary andor at the tonoplast

Study of relationships of trace element concentrations in soilsand plants and their accumulation kinetics are crucial to designphytoextraction applications that maximise the concentrationof a target element in plant biomass Studies in New Caledoniafound significant effects of leaf age on Ni and Mn accumulationwhich could have implications for producing metal-richbiomass to convert into lsquoecocatalystsrsquo (Losfeld et al 2014a)The concept of lsquoecocatalystsrsquo is based on preparation of novelcatalysts from hyperaccumulator biomass These catalystscan be used in the synthesis of molecules using what hasbeen called lsquogreen chemistryrsquo (Losfeld et al 2014b) in whichthese catalysts replace traditional ones The implementationof lsquoecocatalystsrsquo in the organic chemical industry is a strongencouragement for the development of phytoextractionoperations (Grison and Biton 2014) In New Caledoniaphytoextraction with Geissois pruinosa (Cunoniaceae) a Nihyperaccumulator and two subspecies of Grevillea exul(Proteaceae) both Mn accumulators is underway on minewastes (Losfeld et al 2014c) Ongoing large-scale Niphytomining trials in Albania were reported to yield 100 kgNi handash1 from which a high-value product (Ni ammoniumsulfate hexahydrate) was produced (Zhang et al 2014) Usingtwo native Albanian Ni hyperaccumulators (Alyssum muraleand A markgrafii Brassicaceae) a field cropping regimewas developed that yielded up to 17 t handash1 biomass containing120 kg handash1 Ni (Bani et al 2015) The effects of speciescomposition (mono-cropping or co-cropping) with single andmixtures of Ni hyperaccumulator species (Alyssum muraleNoccaea tymphaea Leptoplax emarginata and Bornmuelleratymphaea all Brassicaceae) were also studied in relation to theefficiency of Ni extraction the results showed that biomass andshoot Ni concentrations (though not significant) of B tymphaeaincreased in the co-cropping system (Rue et al 2015)

Recent discoveries of Ni-hyperaccumulator plants werereported from the Philippines in the islands of PalawanSurigao and Zambales where three species of Phyllanthus(Euphorbiaceae) were found containing gt1 foliar Ni Theyinclude Phyllanthus balgooyi with 7638mg gndash1 Ni andP erythrotrichus and P securinegoides both with gt10 000mggndash1 Ni (Quimado et al 2015) Five new Ni hyperaccumulatorswere also reported from Sri Lanka (Euphorbia heterophylla(Euphorbiaceae) Vernonia cinerea (Asteraceae) Flacourtiaindica (Salicaceae) Olax imbricata (Olacaceae) and Toddaliaasiatica (Rutaceae) Iqbal and Rajakaruna 2014) adding to thethree previously documented for the island by Rajakaruna andBohm (2002) The occurrence of copper (Cu) hyperaccumulationamong some ultramafic plants a phenomenon previouslyreported only for five species from Sri Lanka (Rajakaruna andBaker 2004) was also recently confirmed forMalaysia andBrazil(van der Ent and Reeves 2015) Interestingly a laboratory studybyGhasemi et al (2015a) showed that in the presenceofNiCu isaccumulated in both roots and shoots of Ni-hyperaccumulatingAlyssum spp but not in their non-ultramafic congeners Ghasemiet al (2015a) suggested that Cu hyperaccumulation in thepresence of Ni an abundant metal found in serpentine soilsmay be common In Indonesia foliar elemental concentrations

were studied in Emilia sonchifolia (Asteraceae) grown in top andoverburden soils of the local Ni mines (Tjoa and Barus 2014)Their work showed that although this species produced two- tofive-fold greater shoot biomass when grown in top soilsNi-removal rate was higher in overburden soils (which had ahigher Ni concentration) The chromium (Cr) phytoextractionpotential of Brassica juncea (Brassicaceae) was studied inSri Lanka showing that accumulation was dependent on thegenotype and the ionic concentration of the medium highestfoliar accumulation (3511mg gndash1) occurred when plants weresupplied with 200mgmL1 of Cr(VI) (Wijethunga et al 2014)

The use of plants for rehabilitation of mineral wastes wasdemonstrated with Typha angustifolia (Typhaceae) at the formerMamut Copper Mine (Saibeh et al 2014) and by a study onrelationships among soil microbial properties plant cover andbiomass on ultramafic tailings in SouthAfrica (Smith et al 2014)In the latter case marked differences in soil microbial biomassand community structure were observed between the rockdump and the tailings dam with the highest plant biomassbeing recorded at waste piles of intermediate ages and on themoist eastern aspects of the piles The importance of studyingmicrobial relations in phytoremediation efforts was alsoemphasised in a study examining the potential of Hieraciumpilosella (Asteraceae) for phytoremediation in the presence ofsoil microbes (Ogar et al 2014) This study showed beneficialeffects of microbes on plant growth dry mass of shoots androots increased significantly when plants were inoculated withmycorrhizal fungi and nitrogen-fixing bacteria Seneviratne et al(2015) also showed that the inoculation of ultramafic soilswith bacteria and fungi can enhance soil quality and promoteplant growth in the presence of heavy metals Inoculation withmycorrhizal fungi and addition of P can have plant species-specific growth effects as shown by a study investigatingwhether P addition can stimulate plant growth withoutinhibiting mycorrhizal formation (Amir and Gensous 2014)

Geology and soils (Session 2)

Ultramafic rocks containing the serpentine group of mineralsincluding antigorite chrysotile and lizardite have their originswithin the Earthrsquos upper mantle These rocks often form largemassifs and belts or tabular bodies along continental marginsfaults and shear zones Common ultramafic rock types includeperidotites (including dunite wehrlite harzburgite lherzolite)and the secondary alteration products (serpentinites) formedby their hydration within the Earthrsquos crust via a processknown as serpentinisation (Evans et al 2013) Serpentinisationcreates strongly reducing conditions including fluids enrichedwith hydrogen and methane compounds which chemosyntheticmicrobes can exploit for metabolic energy (Cardace and Hoehler2011 McCollom and Seewald 2013 Cardace et al 2014)Detailed accounts of the origins of serpentinites includingtheir mineralogy petrology weathering and geographicdistribution can be found in Coleman and Jove (1992)Moores (2011) and Hirth and Guillot (2013) Over 60 of theglobal Ni supply comes from Ni laterite ores produced from theintensiveweathering of serpentinites found under humid tropicalconditions (Butt andCluzel 2013)Ultramafic soils areweatheredproducts of serpentinite and other ultramafic rocks Ultramafic

4 Australian Journal of Botany A van der Ent et al

bedrock type drainage conditions and weathering intensity allcontribute to differences in pedogenesis and availability ofelements including Ni (Echevarria 2014) For exampleultramafic soils formed by the weathering of peridotite orserpentinite (ultramafic rocks that are chemically similar yetmineralogically distinct) show appreciable differences ingeomorphic and pedologic features (Alexander and DuShey2011) For a summary of the developmental processes ofultramafic soils see Brooks (1987) and Alexander et al(2007) Ultramafic soils are generally deficient in plantessential nutrients such as nitrogen (N) phosphorus (P)potassium (K) and sulfur (S) have a calcium magnesium(Ca Mg) ratio lt1 and have elevated concentrations of heavymetals such as Ni cobalt (Co) Cd and Cr Although physicalfeatures of ultramafic soils can vary considerably from site to site(and within a site) ultramafic soils are often found in open steeplandscapes with substrates that are generally shallow and rockyand that often have reduced capacity for moisture retentionExtensive mining in regions overlying ultramafic bedrockhas led to degraded landscapes needing restoration Carefulmanagement of the topsoil (extracted before the miningdisturbance) can maximise biodiversity on newly restored soilsbecause the topsoil can reintroduce both native seeds andmicrobes to a site under restoration (Bordez et al 2014) Astudy by Mori (2014) examined the potential for greenhousegas emissions (CO2 CH4 N2O) from undisturbed forest soilson different parent materials (sedimentary rock and serpentinerock) along an altitudinal gradient (700 1700 2700 and 3100masl) on Mount Kinabalu Borneo His work suggested thatcarbon fluxes decrease with increasing altitude implying thatlower temperature inhibits microbial activities N2O emissionshowever were not altitude-dependent but were lower onultramafic soils than on sedimentary soils likely because ofthe lower N availability and closed N cycle in ultramafic soilsincluding lower inorganic N nitrification potential microbial Nand higher NH4

+ NO3ndash ratio Whether ultramafic rocks can save

the world from increased carbon emissions has also been a focusof study (Power et al 2013) Mg-rich serpentinite and relatedrocks can react withwater andCO2 to formmagnesium carbonateplus silica thus sequestering potentially damaging CO2

emissions (Maroto-Valer et al 2005 Yang et al 2008) Thistechnique if implemented might have a drastic impact on theunique biota of ultramafic areas

Ecology and biogeography (Session 3)

The unique features of ultramafic habitats can have importanteffects on organismal interactions providing many stimulatingavenues for researchUltramafic sitesmay contain unique speciesof bacteria fungi plants and animals and these often areassembled into communities distinct from surrounding areasFloras of ultramafic sites have attracted considerable attentionat past International Conferences on Serpentine Ecology and the2014Conferencewasnoexception In this session comparisonofultramafic and non-ultramafic forests on Mount Kinabalu (Aibaet al 2015) description of the ultramafic flora of Sri Lanka (Iqbaland Rajakaruna 2014) and a review (Siebert 2014) of SouthAfrican ultramafic-ecology research (which showed that muchwork there has focussed on species descriptions and investigation

of diversity patterns) added to our global knowledge of ultramaficplant ecology Several contributions also added to our floristicknowledge for ultramafic sites in Malaysia (Damit et al 2014Majapun et al 2014 Pereira et al 2014 Sabran et al 2014)including studies conducted at the venue for the poster sessionKinabalu Park (Karim and van der Ent 2014 Sumail and van derEnt 2014) Other presentations contributed floristic informationfrom South Africa (Frisby et al 2014) Japan (Mizuno andKirihata 2015) and Iran (Mohtadi et al 2014) Burgess et al(2015a) documented successional changes that have occurred onultramafic grasslands in the Mid-Atlantic region of the USAshowing that grass-dominated sites have shifted to forests over adecades-long time frame

Rajakaruna and Boyd (2014) pointed out that ultramaficfaunas are little studied compared with ultramafic florasContributions from past International Conferences onSerpentine Ecology (Chazeau 1997 Jourdan 1997 Cantildeameroet al 2004) have added to our knowledge of fauna This traditioncontinued at the 2014 Conference with presentations by Chunget al (2014) on insects from ultramafic sites in a Malaysianforest reserve and Homathevi et al (2014) on termite diversityon ultramafic sites on Mount Tambuyukon Sabah Lichens ofserpentine areas are receiving more study in part because ofwork stimulated bypresentations at past conferences (eg Paukov2009) The 2014 Conference received new contributions on thelichen floras of ultramafic areas in Europe (Paukov et al 2014Favero-Longo et al 2015) and the north-eastern US (Medeiroset al 2014) In contrast to plants and lichens bacteria andmycorrhizal communities seem less affected by ultramaficsoils For example Oline (2006) found little evidence thatthere is a unique ultramafic-soil bacterial flora Venter et al(2015) examined soil algae and cyanoprokaryotes in ultramaficand non-ultramafic soils in South Africa and reached a similarconclusion but did discover that some species were unique to theultramafic communities In a review produced from the 2011conference Southworth et al (2014) found little evidence thatmycorrhizal fungal communities are edaphically specialisedThis may stem from ectomycorrhizal fungi being less inhibitedby edaphic stressors than are plants growing in ultramafic soils(Branco and Ree 2010)

One fascinating component of many ultramafic floras aremetal-hyperaccumulator plants Researchers at the 2014conference continued to catalogue these species describe theirassociates and explore their physiology and ecology van der Entet al (2015) added 19 new Ni-hyperaccumulator species tothe five previously recorded from ultramafic soils in Sabah(Malaysia) illustrating the potential for new discoveries of theseunusual plants Mesjasz-Przybylowicz and Przybylowicz (2014)highlighted intensive and wide-ranging research accomplishedin South Africa on several Ni-hyperaccumulator plant speciesincluding investigations of the mycorrhizal and herbivoreassociates of these plants (also see Mesjasz-Przybyłowicz andPrzybyłowicz 2011) Metal-hyperaccumulator plants may usemetals for defence or the metals may have other functions(such as elemental allelopathy or drought resistance) as recentlyreviewed by Boyd (2014) In a contribution from the 2006conference Boyd (2007) pointed out that most studies of metalhyperaccumulation as a plant defence have focussed on whethermetals increase plant resistance to herbivore attack But tolerance

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 5

of herbivory (ability to withstand damage) may be anotherplant defence tactic affected by metal hyperaccumulationPalomino et al (2007) first addressed this question in a paperpublished in the 2006 conference proceedings findingsignificantly increased tolerance by Noccaea fendleri subspglauca to herbivore damage when plants hyperaccumulated ZnMincey and Boyd (2014) showed elevated tolerance of herbivoryby a Ni-hyperaccumulator species (Streptanthus polygaloidesBrassicaceae) confirming and extending the earlier work with Zn

Finally the patchiness of ultramafic habitats at many sitescan have important impacts on ecological relationships (egpollination seed dispersal) that can affect gene flow andspeciation For example research presented at the 2011conference (Spasojevic et al 2014) explored patterns of seed-dispersal syndromes in Californian ultramafic and non-ultramaficfloras including the influence of ultramafic patch size Substratetype significantly influenced seed dispersal by vertebrates andwind but patch size did not have an important effect At the2014 Conference the patchiness of ultramafic habitats wasevident by presentations from many locations including forexample South Africa Malaysia and the Philippines In thissession Porembski (2014) pointed out similar patchiness ofother geological substrates such as granitoid and ironstoneoutcrops that also results in unique communities Morerecently other edaphic island communities such as those foundon gabbro (Medeiros et al 2015) and gypsum (Moore et al 2014Escudero et al 2015) have also received close attention as settingscomparable to ultramafic habitats for examining ecological andevolutionary theory (Harrison and Rajakaruna 2011) Some of thefloristic works discussed above have important biogeographicaspects because they document the distributions of speciesacross these patchy edaphic landscapes Other contributions tothis session targeted specific biogeographic relationshipsOne such biogeographic concept Rapoportrsquos rule (Veter et al2013) was the topic of the presentation by Whitman and Russo(2014) Rapoportrsquos rule states that the size of a speciesrsquo rangeincreases as altitude or latitude increases andWhitman and Russo(2014) found that the rule applied to soil-generalist species onMount Kinabalu but not to ultramafic-soil specialists Theysuggested that edaphic specialisation limits a species range ofclimatic tolerance (also see Fernandez-Going 2014) A study byGhasemi et al (2015b) examined environmental-stress sensitivityunder different Ca Mg ratios by a ultramafic endemic and itsnon-ultramafic congener Their study suggested that underhigher Ca Mg ratios (characteristic of non-ultramafic soil) theultramafic endemic has reduced tolerance to higher N andincreased temperature The implication of this study is thatultramafic endemics may be more susceptible to human-drivenclimate change-associated stressors such as atmospheric Ndeposition (Pasari et al 2014) than non-ultramafic species Thepresentation by Burge (2014) also addressed the impact ofultramafic soils on plant climatic niches Focusing on soilgeneralists from California USA this study examined whetherultramafic soils allow plants to move outside their climatic nichesThis was indeed the case for both upper and lower limits of speciesdistributions It was suggested that at low elevations ultramaficsoilsprovidea refuge fromcompetitionwhereasathighelevationsthe effect of cold is magnified by infertile ultramafic soils (Burgeand Salk 2014)

Physiology and evolution (Session 4)

Because of the high degree of abiotic stress on ultramafic sitesultramafic-associated biota are model organisms for the study ofadaptation (OrsquoDell and Rajakaruna 2011) The low Ca Mg ratioof ultramafic soils is a major challenge to plant growth andmechanisms to deal with this nutrient imbalance are key toultramafic tolerance These mechanisms include tolerance ofhigh concentrations of soil Mg reduced absorption of Mg orhigher absorption of Ca (Palm and Van Volkenburgh 2014) Inaddition low macronutrient concentrations high concentrationsof some heavy metals and low water availability are alsoimportant factors that must be tolerated by ultramafic plantsSeveral poster and oral presentations at the conference addressedultramafic tolerance with respect to heavy metals and othernutrient imbalances in plants (Doronila et al 2014 Echevarriaet al 2014 Pollard and Smith 2014 Teptina and Paukov 2014Ghaderian et al 2015 Hendry et al 2015) and lichens (Paukovet al 2014 Favero-Longo et al 2015) Although studiesgenerally focus on the effects of a single stress factor thesestress factors are likely to have combined effects (Von Wettberget al 2014) as in the case for Ca Mg ratio and water availabilityon traits of Mimulus guttatus (Phrymaceae Murren et al 2006Selby et al 2014) Presentations also focussed on elementaluptake and localisation in tissues of plants tolerant ofultramafic and other chemically imbalanced soils (Kosugiet al 2015 Mizuno and Kirihata 2015 Pavlova et al 2015)including the use of advanced techniques such as micro-PIXEin mapping regions of elemental concentration within planttissue (Przybyłowicz et al 2005 2014) Tolerance of highconcentrations of heavy metals including molecularmechanisms underlying metal tolerance has also receivedattention (Janssens et al 2009 Gall and Rajakaruna 2013Viehweger 2014) There has been considerable effort tounderstand the linkage between ultramafic-soil stress factorsfor plants (such as metals nutrients water) and the genes thatunderlie adaptations to them (Von Wettberg et al 2014) Recentadvances in molecular biology and genomics have providedpowerful tools to examine the genetic bases for adaptation toultramafic soils (Von Wettberg and Wright 2011 Selby et al2014) In Arabidopsis for example polymorphisms for traitsinvolved in Ca Mg tolerance and heavy-metal detoxificationhave been detected in ultramafic and non-ultramafic populations(Turner et al 2008 2010) suggesting that parallel ecologicaladaptations (Rajakaruna et al 2003 Ostevik et al 2012) canoccur via the differentiation of the same polymorphism atultramafic-tolerant loci in geographically distinct ultramafic-tolerant populations An often used approach for identifyinggenes involved in ultramafic adaptation is the study ofquantitative trait loci (QTL Bratteler et al 2006 Murren et al2006 Wu et al 2008) Recent advances in various fields ofmolecular biology may allow for even newer approaches tostudying ultramafic-plant physiology and the underlyinggenetics of adaptations (Von Wettberg and Wright 2011Selby et al 2014)

Ultramafic plants and their associated biota are also idealmodel systems to explore the role of divergent natural selectionin speciation (Rajakaruna 2004 Kay et al 2011) Intraspecificvariation is central to divergence Population-level variation for

6 Australian Journal of Botany A van der Ent et al

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

bedrock type drainage conditions and weathering intensity allcontribute to differences in pedogenesis and availability ofelements including Ni (Echevarria 2014) For exampleultramafic soils formed by the weathering of peridotite orserpentinite (ultramafic rocks that are chemically similar yetmineralogically distinct) show appreciable differences ingeomorphic and pedologic features (Alexander and DuShey2011) For a summary of the developmental processes ofultramafic soils see Brooks (1987) and Alexander et al(2007) Ultramafic soils are generally deficient in plantessential nutrients such as nitrogen (N) phosphorus (P)potassium (K) and sulfur (S) have a calcium magnesium(Ca Mg) ratio lt1 and have elevated concentrations of heavymetals such as Ni cobalt (Co) Cd and Cr Although physicalfeatures of ultramafic soils can vary considerably from site to site(and within a site) ultramafic soils are often found in open steeplandscapes with substrates that are generally shallow and rockyand that often have reduced capacity for moisture retentionExtensive mining in regions overlying ultramafic bedrockhas led to degraded landscapes needing restoration Carefulmanagement of the topsoil (extracted before the miningdisturbance) can maximise biodiversity on newly restored soilsbecause the topsoil can reintroduce both native seeds andmicrobes to a site under restoration (Bordez et al 2014) Astudy by Mori (2014) examined the potential for greenhousegas emissions (CO2 CH4 N2O) from undisturbed forest soilson different parent materials (sedimentary rock and serpentinerock) along an altitudinal gradient (700 1700 2700 and 3100masl) on Mount Kinabalu Borneo His work suggested thatcarbon fluxes decrease with increasing altitude implying thatlower temperature inhibits microbial activities N2O emissionshowever were not altitude-dependent but were lower onultramafic soils than on sedimentary soils likely because ofthe lower N availability and closed N cycle in ultramafic soilsincluding lower inorganic N nitrification potential microbial Nand higher NH4

+ NO3ndash ratio Whether ultramafic rocks can save

the world from increased carbon emissions has also been a focusof study (Power et al 2013) Mg-rich serpentinite and relatedrocks can react withwater andCO2 to formmagnesium carbonateplus silica thus sequestering potentially damaging CO2

emissions (Maroto-Valer et al 2005 Yang et al 2008) Thistechnique if implemented might have a drastic impact on theunique biota of ultramafic areas

Ecology and biogeography (Session 3)

The unique features of ultramafic habitats can have importanteffects on organismal interactions providing many stimulatingavenues for researchUltramafic sitesmay contain unique speciesof bacteria fungi plants and animals and these often areassembled into communities distinct from surrounding areasFloras of ultramafic sites have attracted considerable attentionat past International Conferences on Serpentine Ecology and the2014Conferencewasnoexception In this session comparisonofultramafic and non-ultramafic forests on Mount Kinabalu (Aibaet al 2015) description of the ultramafic flora of Sri Lanka (Iqbaland Rajakaruna 2014) and a review (Siebert 2014) of SouthAfrican ultramafic-ecology research (which showed that muchwork there has focussed on species descriptions and investigation

of diversity patterns) added to our global knowledge of ultramaficplant ecology Several contributions also added to our floristicknowledge for ultramafic sites in Malaysia (Damit et al 2014Majapun et al 2014 Pereira et al 2014 Sabran et al 2014)including studies conducted at the venue for the poster sessionKinabalu Park (Karim and van der Ent 2014 Sumail and van derEnt 2014) Other presentations contributed floristic informationfrom South Africa (Frisby et al 2014) Japan (Mizuno andKirihata 2015) and Iran (Mohtadi et al 2014) Burgess et al(2015a) documented successional changes that have occurred onultramafic grasslands in the Mid-Atlantic region of the USAshowing that grass-dominated sites have shifted to forests over adecades-long time frame

Rajakaruna and Boyd (2014) pointed out that ultramaficfaunas are little studied compared with ultramafic florasContributions from past International Conferences onSerpentine Ecology (Chazeau 1997 Jourdan 1997 Cantildeameroet al 2004) have added to our knowledge of fauna This traditioncontinued at the 2014 Conference with presentations by Chunget al (2014) on insects from ultramafic sites in a Malaysianforest reserve and Homathevi et al (2014) on termite diversityon ultramafic sites on Mount Tambuyukon Sabah Lichens ofserpentine areas are receiving more study in part because ofwork stimulated bypresentations at past conferences (eg Paukov2009) The 2014 Conference received new contributions on thelichen floras of ultramafic areas in Europe (Paukov et al 2014Favero-Longo et al 2015) and the north-eastern US (Medeiroset al 2014) In contrast to plants and lichens bacteria andmycorrhizal communities seem less affected by ultramaficsoils For example Oline (2006) found little evidence thatthere is a unique ultramafic-soil bacterial flora Venter et al(2015) examined soil algae and cyanoprokaryotes in ultramaficand non-ultramafic soils in South Africa and reached a similarconclusion but did discover that some species were unique to theultramafic communities In a review produced from the 2011conference Southworth et al (2014) found little evidence thatmycorrhizal fungal communities are edaphically specialisedThis may stem from ectomycorrhizal fungi being less inhibitedby edaphic stressors than are plants growing in ultramafic soils(Branco and Ree 2010)

One fascinating component of many ultramafic floras aremetal-hyperaccumulator plants Researchers at the 2014conference continued to catalogue these species describe theirassociates and explore their physiology and ecology van der Entet al (2015) added 19 new Ni-hyperaccumulator species tothe five previously recorded from ultramafic soils in Sabah(Malaysia) illustrating the potential for new discoveries of theseunusual plants Mesjasz-Przybylowicz and Przybylowicz (2014)highlighted intensive and wide-ranging research accomplishedin South Africa on several Ni-hyperaccumulator plant speciesincluding investigations of the mycorrhizal and herbivoreassociates of these plants (also see Mesjasz-Przybyłowicz andPrzybyłowicz 2011) Metal-hyperaccumulator plants may usemetals for defence or the metals may have other functions(such as elemental allelopathy or drought resistance) as recentlyreviewed by Boyd (2014) In a contribution from the 2006conference Boyd (2007) pointed out that most studies of metalhyperaccumulation as a plant defence have focussed on whethermetals increase plant resistance to herbivore attack But tolerance

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 5

of herbivory (ability to withstand damage) may be anotherplant defence tactic affected by metal hyperaccumulationPalomino et al (2007) first addressed this question in a paperpublished in the 2006 conference proceedings findingsignificantly increased tolerance by Noccaea fendleri subspglauca to herbivore damage when plants hyperaccumulated ZnMincey and Boyd (2014) showed elevated tolerance of herbivoryby a Ni-hyperaccumulator species (Streptanthus polygaloidesBrassicaceae) confirming and extending the earlier work with Zn

Finally the patchiness of ultramafic habitats at many sitescan have important impacts on ecological relationships (egpollination seed dispersal) that can affect gene flow andspeciation For example research presented at the 2011conference (Spasojevic et al 2014) explored patterns of seed-dispersal syndromes in Californian ultramafic and non-ultramaficfloras including the influence of ultramafic patch size Substratetype significantly influenced seed dispersal by vertebrates andwind but patch size did not have an important effect At the2014 Conference the patchiness of ultramafic habitats wasevident by presentations from many locations including forexample South Africa Malaysia and the Philippines In thissession Porembski (2014) pointed out similar patchiness ofother geological substrates such as granitoid and ironstoneoutcrops that also results in unique communities Morerecently other edaphic island communities such as those foundon gabbro (Medeiros et al 2015) and gypsum (Moore et al 2014Escudero et al 2015) have also received close attention as settingscomparable to ultramafic habitats for examining ecological andevolutionary theory (Harrison and Rajakaruna 2011) Some of thefloristic works discussed above have important biogeographicaspects because they document the distributions of speciesacross these patchy edaphic landscapes Other contributions tothis session targeted specific biogeographic relationshipsOne such biogeographic concept Rapoportrsquos rule (Veter et al2013) was the topic of the presentation by Whitman and Russo(2014) Rapoportrsquos rule states that the size of a speciesrsquo rangeincreases as altitude or latitude increases andWhitman and Russo(2014) found that the rule applied to soil-generalist species onMount Kinabalu but not to ultramafic-soil specialists Theysuggested that edaphic specialisation limits a species range ofclimatic tolerance (also see Fernandez-Going 2014) A study byGhasemi et al (2015b) examined environmental-stress sensitivityunder different Ca Mg ratios by a ultramafic endemic and itsnon-ultramafic congener Their study suggested that underhigher Ca Mg ratios (characteristic of non-ultramafic soil) theultramafic endemic has reduced tolerance to higher N andincreased temperature The implication of this study is thatultramafic endemics may be more susceptible to human-drivenclimate change-associated stressors such as atmospheric Ndeposition (Pasari et al 2014) than non-ultramafic species Thepresentation by Burge (2014) also addressed the impact ofultramafic soils on plant climatic niches Focusing on soilgeneralists from California USA this study examined whetherultramafic soils allow plants to move outside their climatic nichesThis was indeed the case for both upper and lower limits of speciesdistributions It was suggested that at low elevations ultramaficsoilsprovidea refuge fromcompetitionwhereasathighelevationsthe effect of cold is magnified by infertile ultramafic soils (Burgeand Salk 2014)

Physiology and evolution (Session 4)

Because of the high degree of abiotic stress on ultramafic sitesultramafic-associated biota are model organisms for the study ofadaptation (OrsquoDell and Rajakaruna 2011) The low Ca Mg ratioof ultramafic soils is a major challenge to plant growth andmechanisms to deal with this nutrient imbalance are key toultramafic tolerance These mechanisms include tolerance ofhigh concentrations of soil Mg reduced absorption of Mg orhigher absorption of Ca (Palm and Van Volkenburgh 2014) Inaddition low macronutrient concentrations high concentrationsof some heavy metals and low water availability are alsoimportant factors that must be tolerated by ultramafic plantsSeveral poster and oral presentations at the conference addressedultramafic tolerance with respect to heavy metals and othernutrient imbalances in plants (Doronila et al 2014 Echevarriaet al 2014 Pollard and Smith 2014 Teptina and Paukov 2014Ghaderian et al 2015 Hendry et al 2015) and lichens (Paukovet al 2014 Favero-Longo et al 2015) Although studiesgenerally focus on the effects of a single stress factor thesestress factors are likely to have combined effects (Von Wettberget al 2014) as in the case for Ca Mg ratio and water availabilityon traits of Mimulus guttatus (Phrymaceae Murren et al 2006Selby et al 2014) Presentations also focussed on elementaluptake and localisation in tissues of plants tolerant ofultramafic and other chemically imbalanced soils (Kosugiet al 2015 Mizuno and Kirihata 2015 Pavlova et al 2015)including the use of advanced techniques such as micro-PIXEin mapping regions of elemental concentration within planttissue (Przybyłowicz et al 2005 2014) Tolerance of highconcentrations of heavy metals including molecularmechanisms underlying metal tolerance has also receivedattention (Janssens et al 2009 Gall and Rajakaruna 2013Viehweger 2014) There has been considerable effort tounderstand the linkage between ultramafic-soil stress factorsfor plants (such as metals nutrients water) and the genes thatunderlie adaptations to them (Von Wettberg et al 2014) Recentadvances in molecular biology and genomics have providedpowerful tools to examine the genetic bases for adaptation toultramafic soils (Von Wettberg and Wright 2011 Selby et al2014) In Arabidopsis for example polymorphisms for traitsinvolved in Ca Mg tolerance and heavy-metal detoxificationhave been detected in ultramafic and non-ultramafic populations(Turner et al 2008 2010) suggesting that parallel ecologicaladaptations (Rajakaruna et al 2003 Ostevik et al 2012) canoccur via the differentiation of the same polymorphism atultramafic-tolerant loci in geographically distinct ultramafic-tolerant populations An often used approach for identifyinggenes involved in ultramafic adaptation is the study ofquantitative trait loci (QTL Bratteler et al 2006 Murren et al2006 Wu et al 2008) Recent advances in various fields ofmolecular biology may allow for even newer approaches tostudying ultramafic-plant physiology and the underlyinggenetics of adaptations (Von Wettberg and Wright 2011Selby et al 2014)

Ultramafic plants and their associated biota are also idealmodel systems to explore the role of divergent natural selectionin speciation (Rajakaruna 2004 Kay et al 2011) Intraspecificvariation is central to divergence Population-level variation for

6 Australian Journal of Botany A van der Ent et al

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

of herbivory (ability to withstand damage) may be anotherplant defence tactic affected by metal hyperaccumulationPalomino et al (2007) first addressed this question in a paperpublished in the 2006 conference proceedings findingsignificantly increased tolerance by Noccaea fendleri subspglauca to herbivore damage when plants hyperaccumulated ZnMincey and Boyd (2014) showed elevated tolerance of herbivoryby a Ni-hyperaccumulator species (Streptanthus polygaloidesBrassicaceae) confirming and extending the earlier work with Zn

Finally the patchiness of ultramafic habitats at many sitescan have important impacts on ecological relationships (egpollination seed dispersal) that can affect gene flow andspeciation For example research presented at the 2011conference (Spasojevic et al 2014) explored patterns of seed-dispersal syndromes in Californian ultramafic and non-ultramaficfloras including the influence of ultramafic patch size Substratetype significantly influenced seed dispersal by vertebrates andwind but patch size did not have an important effect At the2014 Conference the patchiness of ultramafic habitats wasevident by presentations from many locations including forexample South Africa Malaysia and the Philippines In thissession Porembski (2014) pointed out similar patchiness ofother geological substrates such as granitoid and ironstoneoutcrops that also results in unique communities Morerecently other edaphic island communities such as those foundon gabbro (Medeiros et al 2015) and gypsum (Moore et al 2014Escudero et al 2015) have also received close attention as settingscomparable to ultramafic habitats for examining ecological andevolutionary theory (Harrison and Rajakaruna 2011) Some of thefloristic works discussed above have important biogeographicaspects because they document the distributions of speciesacross these patchy edaphic landscapes Other contributions tothis session targeted specific biogeographic relationshipsOne such biogeographic concept Rapoportrsquos rule (Veter et al2013) was the topic of the presentation by Whitman and Russo(2014) Rapoportrsquos rule states that the size of a speciesrsquo rangeincreases as altitude or latitude increases andWhitman and Russo(2014) found that the rule applied to soil-generalist species onMount Kinabalu but not to ultramafic-soil specialists Theysuggested that edaphic specialisation limits a species range ofclimatic tolerance (also see Fernandez-Going 2014) A study byGhasemi et al (2015b) examined environmental-stress sensitivityunder different Ca Mg ratios by a ultramafic endemic and itsnon-ultramafic congener Their study suggested that underhigher Ca Mg ratios (characteristic of non-ultramafic soil) theultramafic endemic has reduced tolerance to higher N andincreased temperature The implication of this study is thatultramafic endemics may be more susceptible to human-drivenclimate change-associated stressors such as atmospheric Ndeposition (Pasari et al 2014) than non-ultramafic species Thepresentation by Burge (2014) also addressed the impact ofultramafic soils on plant climatic niches Focusing on soilgeneralists from California USA this study examined whetherultramafic soils allow plants to move outside their climatic nichesThis was indeed the case for both upper and lower limits of speciesdistributions It was suggested that at low elevations ultramaficsoilsprovidea refuge fromcompetitionwhereasathighelevationsthe effect of cold is magnified by infertile ultramafic soils (Burgeand Salk 2014)

Physiology and evolution (Session 4)

Because of the high degree of abiotic stress on ultramafic sitesultramafic-associated biota are model organisms for the study ofadaptation (OrsquoDell and Rajakaruna 2011) The low Ca Mg ratioof ultramafic soils is a major challenge to plant growth andmechanisms to deal with this nutrient imbalance are key toultramafic tolerance These mechanisms include tolerance ofhigh concentrations of soil Mg reduced absorption of Mg orhigher absorption of Ca (Palm and Van Volkenburgh 2014) Inaddition low macronutrient concentrations high concentrationsof some heavy metals and low water availability are alsoimportant factors that must be tolerated by ultramafic plantsSeveral poster and oral presentations at the conference addressedultramafic tolerance with respect to heavy metals and othernutrient imbalances in plants (Doronila et al 2014 Echevarriaet al 2014 Pollard and Smith 2014 Teptina and Paukov 2014Ghaderian et al 2015 Hendry et al 2015) and lichens (Paukovet al 2014 Favero-Longo et al 2015) Although studiesgenerally focus on the effects of a single stress factor thesestress factors are likely to have combined effects (Von Wettberget al 2014) as in the case for Ca Mg ratio and water availabilityon traits of Mimulus guttatus (Phrymaceae Murren et al 2006Selby et al 2014) Presentations also focussed on elementaluptake and localisation in tissues of plants tolerant ofultramafic and other chemically imbalanced soils (Kosugiet al 2015 Mizuno and Kirihata 2015 Pavlova et al 2015)including the use of advanced techniques such as micro-PIXEin mapping regions of elemental concentration within planttissue (Przybyłowicz et al 2005 2014) Tolerance of highconcentrations of heavy metals including molecularmechanisms underlying metal tolerance has also receivedattention (Janssens et al 2009 Gall and Rajakaruna 2013Viehweger 2014) There has been considerable effort tounderstand the linkage between ultramafic-soil stress factorsfor plants (such as metals nutrients water) and the genes thatunderlie adaptations to them (Von Wettberg et al 2014) Recentadvances in molecular biology and genomics have providedpowerful tools to examine the genetic bases for adaptation toultramafic soils (Von Wettberg and Wright 2011 Selby et al2014) In Arabidopsis for example polymorphisms for traitsinvolved in Ca Mg tolerance and heavy-metal detoxificationhave been detected in ultramafic and non-ultramafic populations(Turner et al 2008 2010) suggesting that parallel ecologicaladaptations (Rajakaruna et al 2003 Ostevik et al 2012) canoccur via the differentiation of the same polymorphism atultramafic-tolerant loci in geographically distinct ultramafic-tolerant populations An often used approach for identifyinggenes involved in ultramafic adaptation is the study ofquantitative trait loci (QTL Bratteler et al 2006 Murren et al2006 Wu et al 2008) Recent advances in various fields ofmolecular biology may allow for even newer approaches tostudying ultramafic-plant physiology and the underlyinggenetics of adaptations (Von Wettberg and Wright 2011Selby et al 2014)

Ultramafic plants and their associated biota are also idealmodel systems to explore the role of divergent natural selectionin speciation (Rajakaruna 2004 Kay et al 2011) Intraspecificvariation is central to divergence Population-level variation for

6 Australian Journal of Botany A van der Ent et al

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

adaptive traits and traits that are responsible for reproductiveisolation is known to occur within ultramafic-tolerant speciesacross distinct plant families (OrsquoDell and Rajakaruna 2011)Such intraspecific variation is highlighted in several studies inthis Special Issue including those by Burgess et al (2015b)Chathuranga et al (2015) McAlister et al (2015) and Reeveset al (2015) Research on ultramafic plants including MimulusLayia (Asteraceae) Collinsia (Plantaginaceae) Helianthus(Asteraceae) Noccaea and Lasthenia (Asteraceae) havecontributed greatly to our understanding of factors andmechanisms underlying speciation (Kay et al 2011)demonstrating how edaphic specialisation can greatly reducegene flow among divergent populations (via both pre- and post-zygotic reproductive barriers) setting the stage for subsequentspeciation Ecological approaches including reciprocal-transplantand common-garden experiments have often been employed toexamine local adaptation (Bieger et al 2014) and how ecologicaldivergence can lead to reproductive isolation between closelyrelated taxa (Wright and Stanton 2011 Yost et al 2012) Moyleet al (2012) showed that post-zygotic isolation via hybrid sterilitycan occur between adjacent ultramafic and non-ultramaficecotypes contributing to reduced gene flow and setting thestage for further genetic differentiation via edaphic specialisation

Whether there is a cost associated with ultramafic tolerancehas also been of interest in recent years Studies suggest thatultramafic-tolerant plants are generally less competitive (seeAnacker 2014) and more susceptible to herbivore or pathogenpressure (Lau et al 2008)when found on non-ultramafic soils andthat these biotic factors may drive edaphic specialisationWhether the cost associated with specialisation is greater forendemics than for those only tolerant of ultramafic soil has not yetbeen tested using closely related species pairs or conspecificpopulations (Kay et al 2011) Such studies could shed light onwhy some taxa become endemic to ultramafic soils whereasothers are found on and off ultramafics seemingly indifferent tothe lsquoharshrsquo substrate Themodes of origin for ultramafic endemicsare also of interest (Harris andRajakaruna 2009Kay et al 2011)Phylogenetic studies have suggested that in some cases theevolution of ultramafic endemism is rapid and local (formingneoendemic species Anacker and Strauss 2014) and in othersendemism is via biotype depletion (forming paleoendemicspecies Mayer et al 1994) Anacker et al (2011) andAnacker (2011) have undertaken meta-analyses of molecularphylogenies to explore patterns of diversification on serpentineand the effects of evolutionary and biogeographic histories andregional environmental conditions on the origins of ultramaficendemism Those analyses have shown that ultramafic endemicsexhibit few transitions out of the endemic state suggestingadaptation to ultramafic and subsequent edaphic restriction canlead to an evolutionary lsquodead endrsquo Research by Kolaacuter et al(2012) and Ivaluacute Cacho et al (2014) however suggested thatultramafic lineages may not always represent evolutionary lsquodeadendsrsquo but rather dynamic systems with a potential to furtherdiversify via independent polyploidisation and hybridisationeven providing pathways to radiate off ultramafic soils

Threats and conservation (Session 5)

Inmanyultramafic ecosystemsworldwidemining forNi andothermineral resources contained within ultramafic regoliths presents a

major threat to their biodiversityNowhere is this threat so dire as inNew Caledonia which is one of the worldrsquos most importantbiodiversity hotspots (Wulff et al 2013) The mining industry israpidly growing in New Caledonia with the total production ofNi anticipated to triple in 2015 (Fogliani et al 2014) The NewCaledonian Agronomic Institute conducts research to assist inthe protection of endangered species specifically by identifyingpriority local hotspots and species As an example rare speciessuch as Callitris sulcata (Cupressaceae) and Araucaria rulei(Araucariaceae) are the subject of ongoing population geneticand ecological studies so as to formulate effective species-management plans (Fogliani et al 2014) Elsewhere in thePhilippines the impact of Ni mining is minimised byestablishing vegetation strips in the mined landscape that act asa barrier against soil erosion as well as providing ecosystemcorridors as work in the Caraga Region demonstrates (Varelaet al 2014) These have been termed lsquoecobeltsrsquo and aim tocapture resident biodiversity in degraded areas to assist naturalregeneration over time In Brazil iron ore mining threatensthe unique vegetation of the lsquoIronstone Outcropsrsquo and anenvironmental-impact assessment recently concluded that foursites are now Critically Endangered 11 are Endangered 18are Vulnerable and only one is Stable (Jacobi et al 2014) Thenegative impacts are anticipated to be even higher in thecoming years as a result of habitat loss associated with opencastmining and the lack of officially protected areas

In Sabah where the 8th ICSE was held no active mining formineral resources is taking place at presentHowever the greatestimpact on ultramafic ecosystems results from forest clearing forthe timber industry and for the establishment of palm oilplantations In Sabah 395 of the total forest area existingin 1973 had become deforested by 2010 (Gaveau et al 2014)and protected areas amount to 8 of the land surface (Bryanet al 2013) There are however several large protected areasthat encapsulate important ultramafic ecosystems These includeParks (managed by Sabah Parks) and Forest Reserves (managedby the Forestry Department)

In the United Sates urban development in the San FranciscoBay Area also threatens the long-term viability of ultramaficecosystems (US Fish and Wildlife Service 1998) Regulatorytools are being used there to balance urban development withconservation of the regionrsquos unique ultramafic-associated floraand fauna (Harris 2014) Detailed studies on the genetics andhabitat requirements of coyote ceanothus (Ceanothus ferrisiaeRhamnaceae) a rare shrub have led to the identification of asuitable introduction site to mitigate the impending threats to itslargest population as a result of planned dam constructionactivities (Hillman et al 2014)

Although not of ultramafic origin the CopperndashCobalt Belt ofthe Democratic Republic of the Congo and Zambia is the richestglobally formetallophytes and hyperaccumulators with over 600such species having been described (Malaisse 2014) Researchundertaken by the universities of Lubumbashi Liegravege andBrussels has led to the creation of an online database (wwwcopperfloraorg accessed 31 March 2015) and to an upcomingbook entitled lsquoCopper-cobalt flora of Katanga and northernZambiarsquo Unfortunately this unique ecosystem is under acutethreat because many of the lsquocopper hillsrsquo are being mined andseveral species are already thought extinct (Malaisse 2014)

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 7

Although the rich biodiversity of many ultramafic ecosystemsis often emphasised limited research on these ecosystems hasbeen undertaken in some of the most biodiverse regions of theworld For example the ultramafic outcrops on the islands ofSulawesi and Halmahera in Indonesia covering 15 400 km2 andgt5500 km2 respectively have hardly been studied (van der Entet al 2013b) Borneo Island has approximately 14 500 speciesof plants (Roos et al 2004) and at least 4000 of these occuron ultramafic soils (van der Ent et al 2015) Without adequateinformation it is difficult to formulate conservation strategiesand priorities although given the ongoing habitat destructionit is evident that many plant species are threatened Anothervery specific threat to ultramafic ecosystems is brought about bylateritic Ni-mining activitiesmainly in Indonesia the Philippinesand New Caledonia These mining activities target Ni-richultramafic soils which often host species-rich vegetation witha high percentage of endemic species Species adapted to thriveon ultramafic soils offer rich genetic resources for mine-siterehabilitation after strip-mining but are likely to be destroyedduring the mining operations

These global examples have highlighted that ultramaficecosystems are under threat in many areas around the worldThe combined forces of urbanisation mineral extraction andconversion of forests to plantations cast a bleak forecast forthe survival of many rare and endemic plant and other speciesAlthough not frequently emphasised the lack of scientificknowledge of the plant diversity and ecology of ultramaficoutcrops in South-east Asia and other less explored parts ofthe world puts constraints on protecting this biodiversity withmany species having already disappeared unnoticed Thisexemplifies the urgent need for more research to assist in theconservation of areasmost under threat (Whiting et al 2004) Thepotential effects of climate change may have a further impact onthe survival of many species restricted to ultramafic soils (seeFernandez 2014 and references cited therein)

Diversity systematics and taxonomy (Session 6)

Ultramafic outcrops are home to unique plant communitiescharacterised by rare and endemic species (Kruckeberg 1984Brooks 1987 Alexander et al 2007 Rajakaruna et al 2009 vander Ent et al 2014b) Ultramafic outcrops are extensive in Sabahcovering an area of ~3500 km2 and occur from sea level up tonearly 3000m asl (Proctor et al 1988 Repin 1998) Sabah isextremely biodiverse with an estimated 8000 plant species(Wong 1992) of which at least 800 species are endemic(Maycock et al 2012) In total 4252 species have beenrecorded from the ultramafic soils in Sabah (van der Ent et al2014a) Within Sabah Kinabalu Park is the pinnacle of plantdiversity with over 5000 plant species in an area lt1200 km2 themost species-rich area on Earth in terms of species density(Beaman 2005) In Kinabalu Park ultramafic soils support atleast 2854 plant species in 742 genera and 188 families (van derEnt et al 2014a) Elsewhere in Sabah there is remarkablediversity of insects in the ultramafic Tinkar Forest Reserve(Chung et al 2014) Also in the Sungai Imbak Virgin JungleReserve 62 Dipterocarpaceae species from six genera wererecorded including 15 species as new records (Majapun et al2014) The biodiversity riches of ultramafic outcrops at Tawau

Forest Reserve and in the Mount Silam Forest Reserve were alsonoted during the conference (Nilus et al 2014 Pereira et al2014) The latter contained 282 taxa from 72 families some arenarrow endemics known only from that site Elsewhere in Asiaremote-sensing studieswere undertaken in theAndamangroupofislands of India where supervised classification of the vegetationrevealed three distinct outcrops in Rutland Island and on the hilltop of Chidyatappu Island (Datta et al 2014) and a dwarf scrubcommunity on the top of the Saddle Hills in North AndamanIsland (Chaudhury et al 2014)

Taxonomic studies on Cleisocentron (Orchidaceae) andTernstroemia (Pentaphylacaceae) revealed several speciesrestricted to ultramafic soils in Sabah (Nyee-Fan et al 2014Sabran et al 2014) Species of carnivorous pitcher plantsin Nepenthes have attracted significant interest in Sabah(Clarke 1997) with 22 of the 39 Bornean Nepenthes found inSabah (Damit et al 2014) Kinabalu Park is themost species-rich(14 species) five species are endemic to Kinabalu Park namelyN burbidgeaeN edwardsianaN rajah andN villosa Detailedecological studies in Kinabalu Park have shown thatphysiognomy rather than co-occurring species compositiondetermines habitat differentiation of Nepenthes species (Sumailand Van der Ent 2014)

Elsewhere on the globe the plant diversity of ironstoneoutcrops in south-eastern Brazil was highlighted by Jacobiet al (2014) Also in South Africa in the vegetation ofGriqualand most endemic plants were edaphic specialistswith a preference for specific geologic substrates howeverdistribution modelling indicated that climate overrides edaphicpreference in some instances (Frisby et al 2014) Anotherstudy from South Africa focused on plantndashsoil relations ofthe Vredefort Dome (an impact structure) and how abrupttransitions in soil chemical characteristics can ultimatelyaffect the floristic and physiognomic characteristics of theassociated vegetation (Boneschans et al 2015) Finallyecological succession on Cu-contaminated mine tailings in thePhilippines was reported to consist of Digitaria sanguinalisPaspalum conjugatum and P scrobiculum (all Poaceae) in theearly stages which are then replaced byP conjugatumCynodondactylon (Poaceae) and Cuphea carthagenensis (Lythraceae)with associated increases in soil pH and organic matter content(Cuevas and Balangcod 2014)

Summary

Ultramafic geoecology is now a significant focus in the naturalsciences A recent (20 February 2015) search on Web of Science(from all databases) of the topic lsquoserpentine and ecologyrsquo yielded853 results with 8688 citing articles (without self-citations)over the past 20 years with an exponential growth in citationsduring the past decade Studies conducted in geology pedologymicrobiology ecology evolution conservation and restoration ofultramafic ecosystems continue to unearth unresolved questionssetting thestage formultidisciplinaryand interdisciplinary researchworldwide We have summarised some of the contributionsmade during the 8th International Conference on SerpentineEcology held in Sabah Malaysia and described some of theexciting challenges awaiting future research in this double issue(1ndash2 combined) and the next double issue (3ndash4 combined) of

8 Australian Journal of Botany A van der Ent et al

Australian Journal of Botany We hope that the 9th InternationalConference (scheduled for 2017 and hosted by The AgriculturalUniversity of Tirana in Albania) will be as successful as werethe 8th (and prior) conferences in advancing our knowledge ofultramafic ecosystems worldwide

Acknowledgements

We are grateful to the Director of Sabah Parks Dr Jamili Nais to Chairman ofSabahParksBoardofTrusteesDatorsquoSeriTengkuDrZainalAdlinBinTengkuMahamood and to the Minister for Tourism Culture and EnvironmentYB Datuk Seri Panglima Masidi Manjun for their support to host the8th International Serpentine Ecology Conference in Sabah (Malaysia)Additionally we acknowledge the many reviewers for their expert advicethat greatly improved the quality of the manuscripts presented in this SpecialIssue Robert Boyd acknowledges support from the Alabama AgriculturalExperiment Station Project No ALA021-1-09008

References

Aiba S SawadaY TakyuM SeinoTKitayamaKRepinR (2015) Structurefloristics and diversity of tropical montane rain forests over ultramaficsoils onMountKinabalu (Borneo) comparedwith those onnon-ultramaficsoils Australian Journal of Botany 63 doi101071BT14238

Alexander EB DuShey J (2011) Topographic and soil differences fromperidotite to serpentinite In lsquoSpecial issue driving forces for globalpedodiversityrsquo Geomorphology 135 271ndash276doi101016jgeomorph201102007

Alexander EB ColemanRB Keeler-Wolf T Harrison SP (2007) lsquoSerpentinegeoecology of western North Americarsquo (Oxford University Press NewYork)

Amir H Gensous S (2014) Combined effects of arbuscular mycorrhizalcolonization and phosphorus concentration level on growth andadaptation of three endemic plant species to ultramafic soil In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd SStrauss NRRajakarunaAJPollardMLakimAvanderEntMSuleiman JBSugau)pp 39ndash39 (SabahParksKota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Anacker BL (2011) Phylogenetic patterns of endemism and diversity InlsquoSerpentine the evolution and ecology of a model systemrsquo (Eds SPHarrison N Rajakaruna) pp 49ndash79 (University of California PressBerkeley CA)

Anacker BL (2014) The nature of serpentine endemism American Journal ofBotany 101 219ndash224 doi103732ajb1300349

Anacker BL Strauss SY (2014) The geography and ecology of plantspeciation range overlap and niche divergence in sister speciesProceedings of the Royal Society Series B 281 20132980doi101098rspb20132980

Anacker BL Whittall J Goldberg E Harrison SP (2011) Origins andconsequences of serpentine endemism in the California flora Evolution65 365ndash376 doi101111j1558-5646201001114x

Baker AJM (2014) Phytoremediation emerged or still emergingphytotechnology In lsquoOfficial Programme Book (oral presentations)8th International Conference on Serpentine Ecology (9ndash13 June2014)rsquo (Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RSBoyd S Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 21 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Baker AJM Proctor J Reeves RD (Eds) (1992) lsquoThe vegetation of ultramafic(serpentine) soils proceedings of the first international conference onserpentine ecologyrsquo (Intercept Andover UK)

Balkwill K (Ed) (2001) Proceedings third international conference onserpentine ecology Part 2special issue South African Journal ofScience 97

Bani A Echevarria G Zhang X Laubie B Benizri E Morel JL SimonnotM-O (2015) The effect of plant density in nickel phytomining fieldexperiments with Alyssum murale in Albania Australian Journal ofBotany 63 72ndash77 doi101071BT14285

Beaman JH (2005) Mount Kinabalu hotspot of plant diversity in BorneoBiologiske Skrifter 55 103ndash127

Bieger A Rajakaruna N Harrison SP (2014) Little evidence for localadaptation to soils or microclimate in the postfire recruitment of threeCalifornian shrubs Plant Ecology amp Diversity 7 411ndash420doi101080175508742012701670

Boneschans R Coetzee M Siebert SJ (2015) A geobotanical investigationof the Koedoesfontein Complex Vredefort Dome South AfricaAustralian Journal of Botany 63 doi101071BT14267

Boominathan R Saha-Chaudhury NM Sahajwalla V Doran PM (2004)Production of nickel bio-ore from hyperaccumulator plant biomassapplications in phytomining Biotechnology and Bioengineering 86243ndash250 doi101002bit10795

BordezL JourandPDucoussoMCarricondeFCavalocYSantini SMozarMLeveauAAmirH (2014) Selective topsoilmanagement ofmined landfor vegetation communities restoration In lsquoOfficial Programme Book(oral presentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 38 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Borhidi A (1992) The serpentine flora and vegetation of Cuba In lsquoThevegetation of ultramafic (serpentine) soils proceedings of the firstinternational conference on serpentine ecologyrsquo (Eds AJM Baker JProctor RD Reeves) pp 83ndash95 (Intercept Andover UK)

Boyd RS (2007) The defense hypothesis of elemental hyperaccumulationstatus challenges and new directions Plant and Soil 293 153ndash176doi101007s11104-007-9240-6

BoydRS (2014) Ecology and evolution of metal-hyperaccumulator plants InlsquoPlant ecology and evolution in harsh environmentsrsquo (Eds NRajakarunaRS Boyd T Harris) pp 227ndash241 (Nova Science Publishers HauppaugeNY)

Boyd RS Baker AJM Proctor J (Eds) (2004) lsquoUltramafic rocks their soilsvegetation and faunarsquo (Science Reviews 2000 St Albans UK)

Boyd RS Kruckeberg AR Rajakaruna N (2009) Biology of ultramafic rocksand soils research goals for the future Northeastern Naturalist 16422ndash440 doi1016560450160530

Branco S Ree R (2010) Serpentine soils do not limit ectomycorrhizalfungal diversity PLoS ONE 5(7) e11757doi101371journalpone0011757

Bratteler M Lexer C Widmer A (2006) Genetic architecture of traitsassociated with serpentine adaptation of Silene vulgaris Journal ofEvolutionary Biology 19 1149ndash1156doi101111j1420-9101200601090x

Brooks RR (1987) lsquoSerpentine and its vegetation a multidisciplinaryapproachrsquo (Dioscorides Press Portland OR)

Bryan JE Shearman PL Asner GP Knapp DE Aoro G Lokes B (2013)Extreme differences in forest degradation in Borneo comparing practicesin Sarawak Sabah and Brunei PLoS ONE 8(7) e69679doi101371journalpone0069679

Burge DO (2014) How do infertile soils alter the climatic niche of generalistplant species In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJMBaker RDReeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 58 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 9

Australian Journal of Botany We hope that the 9th InternationalConference (scheduled for 2017 and hosted by The AgriculturalUniversity of Tirana in Albania) will be as successful as werethe 8th (and prior) conferences in advancing our knowledge ofultramafic ecosystems worldwide

Acknowledgements

We are grateful to the Director of Sabah Parks Dr Jamili Nais to Chairman ofSabahParksBoardofTrusteesDatorsquoSeriTengkuDrZainalAdlinBinTengkuMahamood and to the Minister for Tourism Culture and EnvironmentYB Datuk Seri Panglima Masidi Manjun for their support to host the8th International Serpentine Ecology Conference in Sabah (Malaysia)Additionally we acknowledge the many reviewers for their expert advicethat greatly improved the quality of the manuscripts presented in this SpecialIssue Robert Boyd acknowledges support from the Alabama AgriculturalExperiment Station Project No ALA021-1-09008

References

Aiba S SawadaY TakyuM SeinoTKitayamaKRepinR (2015) Structurefloristics and diversity of tropical montane rain forests over ultramaficsoils onMountKinabalu (Borneo) comparedwith those onnon-ultramaficsoils Australian Journal of Botany 63 doi101071BT14238

Alexander EB DuShey J (2011) Topographic and soil differences fromperidotite to serpentinite In lsquoSpecial issue driving forces for globalpedodiversityrsquo Geomorphology 135 271ndash276doi101016jgeomorph201102007

Alexander EB ColemanRB Keeler-Wolf T Harrison SP (2007) lsquoSerpentinegeoecology of western North Americarsquo (Oxford University Press NewYork)

Amir H Gensous S (2014) Combined effects of arbuscular mycorrhizalcolonization and phosphorus concentration level on growth andadaptation of three endemic plant species to ultramafic soil In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd SStrauss NRRajakarunaAJPollardMLakimAvanderEntMSuleiman JBSugau)pp 39ndash39 (SabahParksKota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Anacker BL (2011) Phylogenetic patterns of endemism and diversity InlsquoSerpentine the evolution and ecology of a model systemrsquo (Eds SPHarrison N Rajakaruna) pp 49ndash79 (University of California PressBerkeley CA)

Anacker BL (2014) The nature of serpentine endemism American Journal ofBotany 101 219ndash224 doi103732ajb1300349

Anacker BL Strauss SY (2014) The geography and ecology of plantspeciation range overlap and niche divergence in sister speciesProceedings of the Royal Society Series B 281 20132980doi101098rspb20132980

Anacker BL Whittall J Goldberg E Harrison SP (2011) Origins andconsequences of serpentine endemism in the California flora Evolution65 365ndash376 doi101111j1558-5646201001114x

Baker AJM (2014) Phytoremediation emerged or still emergingphytotechnology In lsquoOfficial Programme Book (oral presentations)8th International Conference on Serpentine Ecology (9ndash13 June2014)rsquo (Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RSBoyd S Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 21 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Baker AJM Proctor J Reeves RD (Eds) (1992) lsquoThe vegetation of ultramafic(serpentine) soils proceedings of the first international conference onserpentine ecologyrsquo (Intercept Andover UK)

Balkwill K (Ed) (2001) Proceedings third international conference onserpentine ecology Part 2special issue South African Journal ofScience 97

Bani A Echevarria G Zhang X Laubie B Benizri E Morel JL SimonnotM-O (2015) The effect of plant density in nickel phytomining fieldexperiments with Alyssum murale in Albania Australian Journal ofBotany 63 72ndash77 doi101071BT14285

Beaman JH (2005) Mount Kinabalu hotspot of plant diversity in BorneoBiologiske Skrifter 55 103ndash127

Bieger A Rajakaruna N Harrison SP (2014) Little evidence for localadaptation to soils or microclimate in the postfire recruitment of threeCalifornian shrubs Plant Ecology amp Diversity 7 411ndash420doi101080175508742012701670

Boneschans R Coetzee M Siebert SJ (2015) A geobotanical investigationof the Koedoesfontein Complex Vredefort Dome South AfricaAustralian Journal of Botany 63 doi101071BT14267

Boominathan R Saha-Chaudhury NM Sahajwalla V Doran PM (2004)Production of nickel bio-ore from hyperaccumulator plant biomassapplications in phytomining Biotechnology and Bioengineering 86243ndash250 doi101002bit10795

BordezL JourandPDucoussoMCarricondeFCavalocYSantini SMozarMLeveauAAmirH (2014) Selective topsoilmanagement ofmined landfor vegetation communities restoration In lsquoOfficial Programme Book(oral presentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 38 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Borhidi A (1992) The serpentine flora and vegetation of Cuba In lsquoThevegetation of ultramafic (serpentine) soils proceedings of the firstinternational conference on serpentine ecologyrsquo (Eds AJM Baker JProctor RD Reeves) pp 83ndash95 (Intercept Andover UK)

Boyd RS (2007) The defense hypothesis of elemental hyperaccumulationstatus challenges and new directions Plant and Soil 293 153ndash176doi101007s11104-007-9240-6

BoydRS (2014) Ecology and evolution of metal-hyperaccumulator plants InlsquoPlant ecology and evolution in harsh environmentsrsquo (Eds NRajakarunaRS Boyd T Harris) pp 227ndash241 (Nova Science Publishers HauppaugeNY)

Boyd RS Baker AJM Proctor J (Eds) (2004) lsquoUltramafic rocks their soilsvegetation and faunarsquo (Science Reviews 2000 St Albans UK)

Boyd RS Kruckeberg AR Rajakaruna N (2009) Biology of ultramafic rocksand soils research goals for the future Northeastern Naturalist 16422ndash440 doi1016560450160530

Branco S Ree R (2010) Serpentine soils do not limit ectomycorrhizalfungal diversity PLoS ONE 5(7) e11757doi101371journalpone0011757

Bratteler M Lexer C Widmer A (2006) Genetic architecture of traitsassociated with serpentine adaptation of Silene vulgaris Journal ofEvolutionary Biology 19 1149ndash1156doi101111j1420-9101200601090x

Brooks RR (1987) lsquoSerpentine and its vegetation a multidisciplinaryapproachrsquo (Dioscorides Press Portland OR)

Bryan JE Shearman PL Asner GP Knapp DE Aoro G Lokes B (2013)Extreme differences in forest degradation in Borneo comparing practicesin Sarawak Sabah and Brunei PLoS ONE 8(7) e69679doi101371journalpone0069679

Burge DO (2014) How do infertile soils alter the climatic niche of generalistplant species In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJMBaker RDReeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 58 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 9

Burge DO Salk CF (2014) Climatic niche shifts in the serpentine soil flora ofCalifornia Journal of Vegetation Science 25 873ndash884doi101111jvs12144

Burgess JL Szlavecz K Rajakaruna N Lev S Swan CM (2015a) Vegetationdynamics and mesophication in response to conifer encroachment withinan ultramafic system Australian Journal of Botany 63doi101071BT14241

Burgess JL Szlavecz K Rajakaruna N Swan CM (2015b) Ecotypicdifferentiation of mid-Atlantic Quercus species in response toultramafic soils Australian Journal of Botany 63 doi101071BT14274

Butt CRM Cluzel D (2013) Nickel laterite ore deposits weatheredserpentinites Elements 9 123ndash128 doi102113gselements92123

Cantildeamero AB Aacuteguila RN Rodriacuteguez-Fernaacutendez K (2004) The Lepidopteraof plant formations on Cuban ultramafics a preliminary analysis InlsquoUltramafic rocks their soils vegetation and fauna proceedings of thefourth international conference on serpentine ecology 21ndash26April 2003rsquo(Eds RS Boyd AJM Baker J Proctor) pp 223ndash226 (Science ReviewsSaint Albans UK)

Cardace D Hoehler TM (2011) Microbes in extreme environmentsimplications for life on the early Earth and other planets InlsquoSerpentine the evolution and ecology of a model systemrsquo (Eds SPHarrison N Rajakaruna) pp 29ndash48 (University of California PressBerkeley CA)

Cardace D Meyer-Dombard DR Olsen AA Parenteau MN (2014) Bedrockand geochemical controls on extremophile habitats In lsquoPlant ecology andevolution in harsh environmentrsquosrsquo (EdsNRajakaruna RSBoyd THarris)pp 1ndash32 (Nova Science Publishers Hauppauge NY)

Chaney RL Angle JS Broadhurst CL Peters CA Tappero RV Sparks DL(2007) Improved understanding of hyperaccumulation yields commercialphytoextractionandphytomining technologies JournalofEnvironmentalQuality 36 1429ndash1443 doi102134jeq20060514

Chaney RL Reeves RD Baklanov IA Centofanti T Broadhurst CL BakerAJM Van der Ent A Roseberg RJ (2014) Phytoremediation andphytomining using plants to remediate contaminated or mineralizedenvironments In lsquoPlant ecology and evolution in harsh environmentsrsquo(Eds N Rajakaruna RS Boyd T Harris) pp 365ndash392 (Nova SciencePublishers Hauppauge NY)

Chathuranga PKD Dharmasena SKAT Rajakaruna N Iqbal MCM (2015)Growth and nickel uptake by serpentine and non-serpentine populationsof Fimbristylis ovata (Cyperaceae) from Sri Lanka Australian Journal ofBotany 63 128ndash133 doi101071BT14232

Chaudhury K Dutta S Mukherjee PK (2014) Plant diversity characterisationof serpentine outcrops on Saddle Hills using remote sensing technologyof North Andaman Islands India In lsquoOfficial Programme Book (oralpresentations) 8th InternationalConference onSerpentineEcology (9ndash13June 2014)rsquo (EdsAJMBakerRDReevesGEchevarriaLLrsquoHuillierRSBoyd S Strauss NRRajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 83 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Chazeau J (1997) Caractegraveres de la faune de quelquesmilieux naturels sur solsultramafiques enNouvelle-Caleacutedonie In lsquoEacutecologie desmilieux sur rochesultramafiques et sur sols meacutetallifegraveres actes de la deuxiegraveme confeacuterenceinternationale sur lrsquoecologie des milieux serpentiniques Noumeacutea 31juillet ndash 5 aoucirct 1995rsquo (Eds T Jaffreacute RD Reeves T Becquer)pp 95ndash105 Documents Scientifiques et Techniques III2 (CentreORSTOM de Noumeacutea Noumeacutea New Caledonia)

Chiarucci A Baker AJM (2007) Proceedings of the fifth internationalconference on serpentine ecology Plant and Soil 293

Chung AYC Nurul Aqidah I John LY Tajuddin MA Nilus R (2014)Exploring insects of the ultramafic Gunung Tinkar Forest Reserve incentral Sabah Malaysia In lsquoOfficial Programme Book (oralpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria L

LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 84 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Clarke CM (1997) lsquoNepenthes of Borneorsquo (Natural History Publications(Borneo) Kota Kinabalu Malaysia)

Coleman RG Jove C (1992) Geological origin of serpentinites In lsquoThevegetation of ultramafic (serpentine) soils proceedings of the firstinternational conference on derpentine rcologyrsquo (Eds AJM Baker JProctor RD Reeves) pp 469ndash494 (Intercept Andover UK)

Cuevas VC Balangcod TM (2014) Ecological succession in three copper-contaminated mine tailing ponds in Mankayan Benguet ProvinceCordillera Administrative Region Philippines In lsquoOfficial ProgrammeBook (oral presentations) 8th International Conference on SerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarriaLLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M Lakim Avan der Ent M Suleiman JB Sugau) p 85 (Sabah Parks Kota KinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Damit A Pereira JT Sabran S En CT (2014) The distribution of Nepenthesspecies on ultramafic soils in Sabah Malaysia In lsquoOfficial ProgrammeBook (poster presentations) 8th International Conference on SerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarriaLLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M Lakim Avan der Ent M Suleiman JB Sugau) p 35 (Sabah Parks Kota KinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Datta S ChaudhuryKMukherjee PK (2014) Serpentinophytes fromRutlandandChidyatappu of SouthAndaman India In lsquoOfficial ProgrammeBook(oral presentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 86 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Doronila AI Quimado MO Dias DA Fernando ES (2014) The metaboliteand elemental content of four Ni hyperaccumulators from ZambalesPhilippines In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 33 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Echevarria G (2014) The availability of nickel in ultramafic soils In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd S StraussNRRajakarunaAJ PollardMLakimAvanderEntMSuleiman JBSugau)p 37 (SabahParksKotaKinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Echevarria G Baker AJM Bani A Konstantinou M Zhang X Benizri ESimonnotMOMorel J-L (2014) Hyperaccumulation of Ni in serpentinesof Albania and continental Greece what are the trade-offs betweenaccumulated elements In lsquoOfficial Programme Book (oralpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimA van der Ent M Suleiman JB Sugau) p 64ndash65 (Sabah Parks KotaKinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Escudero A Palacio S Maestre FT Luzuriaga AL (2015) Plant life ongypsum a review of its multiple facets Biological Reviews of theCambridge Philosophical Society 90 1ndash18 doi101111brv12092

10 Australian Journal of Botany A van der Ent et al

Evans BK Hattori K Baronnet A (2013) Serpentinite what why whereElements 9 99ndash106 doi102113gselements9299

Favero-LongoSEMatteucci EMorandoMRolfo FHarris TB Piervittori R(2015) Metals and secondary metabolites in saxicolous lichencommunities on ultramafic and non-ultramafic rocks of the westernItalian Alps Australian Journal of Botany 63 doi101071BT14256

Fernandez-GoingBM (2014)Climate change and the future of edaphicflorasIn lsquoPlant ecology and evolution in harsh environmentsrsquo (Eds NRajakaruna RS Boyd T Harris) pp 297ndash312 (Nova SciencePublishers Hauppauge NY)

Fernando DR Marshall AT Hoebee SE Baker AJM Siegele R Forster PI(2014) Australian Gossia generic manganese accumulators specific toother metals In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 24 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Fogliani B Wulff A Anquez M Haverkamp C LrsquoHuillier L (2014)Biodiversity conservation in New Caledonia IAC programs forendangered plant species and flora conservation on ultramaficenvironment In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 74 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Frisby A Siebert SJ Cilliers D van Wyk AE (2014) Redefining theGriqualand west centre of endemism ndash a third ultramafic flora fromSouth Africa In lsquoOfficial Programme Book (poster presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJMBaker RDReeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 29 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-poster

Gall JE Rajakaruna N (2013) The physiology functional genomics andapplied ecology of heavy metal-tolerant Brassicaceae In lsquoBrassicaceaecharacterization functional genomics and health benefitsrsquo (Ed M Lang)pp 121ndash148 (Nova Science Publishers Hauppauge NY)

Gaveau DLA Sloan S Molidena E Yaen H Sheil D Abram NK AncrenazM Nasi R Quinones M Wielaard N Meijaard E (2014) Four decades offorest persistence clearance and logging on Borneo PLoS ONE 9(7)e101654 doi101371journalpone0101654

Ghaderian SM Ghasemi R Hajihashemi F (2015) Interaction of nickeland manganese in uptake translocation and accumulation by thenickel hyperaccumulator plant Alyssum bracteatum (Brassicaceae)Australian Journal of Botany 63 47ndash55 doi101071BT14210

Ghasemi R Ghaderian SM Ebrazeh S (2015a) Nickel stimulates copperuptake by nickel hyperaccumulator plants in the genus AlyssumAustralian Journal of Botany 63 56ndash64 doi101071BT14219

Ghasemi R Zare Chavoshi Z Boyd RS Rajakaruna N (2015b)Calcium magnesium ratio affects environmental stress sensitivity inthe serpentine-endemic Alyssum inflatum (Brassicaceae) AustralianJournal of Botany 63 39ndash46 doi101071BT14235

Grison C Biton J (2014) Phytoextraction and ecological catalysis symbiosisfor future In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) pp 34ndash35 (Sabah Parks Kota KinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Guillot S Hattori K (2013) Serpentinites essential roles in geodynamicsarc volcanism sustainable development and the origin of lifeElements9 95ndash98 doi102113gselements9295

Harris T (2014) Conservation of serpentine-endemic species in the SanFrancisco Bay Area California USA In lsquoOfficial Programme Book(oral presentations) 8th International Conference on SerpentineEcology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria L LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJPollard M Lakim A van der Ent M Suleiman JB Sugau) p 76(Sabah Parks Kota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Harris TB Rajakaruna N (2009)Adiantum viridimontanum Aspidotis densaMinuartia marcescens and Symphyotrichum rhiannon additionalserpentine endemics from eastern North America NortheasternNaturalist 16 111ndash120 doi1016560450160509

Harrison SP Rajakaruna N (Eds) (2011) lsquoSerpentine evolution andecology of a model systemrsquo (University of California Press BerkeleyCA)

HendryRWormingtonKWalshK (2015) An ecological study of the centralQueensland ultramafic endemic shrub Neoroepera buxifoliaMuell Argamp FMuell (Picrodendraceae) Australia Australian Journal of Botany63 doi101071BT14184

Hillman JM Honeycutt RL Vasey MC Berry AM Potter D Gardipee FM(2014) Dam the luck How the Anderson Dam Seismic Retrofit has led tonew insights into the ecology evolution and conservation of the coyoteceanothus (Ceanothus ferrisiae) In lsquoOfficial Programme Book (posterpresentations) 8th International ConferenceonSerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarria LLrsquoHuillierRSBoyd S Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 38 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-poster

Hirth G Guillot S (2013) Rheology and tectonic significance of serpentiniteElements 9 107ndash113 doi102113gselements92107

Homathevi R WongMK HongMC (2014) Termite species diversity withinultramafic area of Mount Tambuyukon Sabah Malaysia In lsquoOfficialProgramme Book (poster presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd SStraussNRRajakarunaAJPollardMLakimAvanderEntMSuleiman JBSugau)p 45 (SabahParksKotaKinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Iqbal MCM Rajakaruna N (2014) Serpentine ecology in Sri Lanka currentknowledge information gaps and future directions Conservation ofserpentine-endemic species in the San Francisco Bay Area CaliforniaUSA In lsquoOfficial ProgrammeBook (oral presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM BakerRD Reeves G Echevarria L LrsquoHuillier RS Boyd S Strauss NRRajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 53 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Ivaluacute CachoNBurrell AM Pepper A Strauss SY (2014) Systematics and theevolution of serpentine tolerance in the California jewelflowers(Streptanthus) and its allies Molecular Phylogenetics and Evolution72 71ndash81

Jacobi CMDoCarmoFFDoCarmoFF (2014) Threats to ironstone outcropsplant communities in southeast Brazil In lsquoOfficial ProgrammeBook (oralpresentations) 8th International ConferenceonSerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarria LLrsquoHuillierRSBoyd S Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 46 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 11

Jaffreacute T (1992) Floristic and ecological diversity of the vegetation onultramafic rocks in New Caledonia In lsquoThe vegetation of ultramafic(serpentine) soils proceedings of the first international conference onserpentine ecologyrsquo (Eds AJM Baker J Proctor RD Reeves)pp 101ndash107 (Intercept Andover UK)

Jaffreacute TReevesRDBecquerT (Eds) (1997) lsquoEacutecologie desmilieux sur rochesultramafiques et sur sols meacutetallifegraveres actes de la deuxiegraveme confeacuterenceinternationale sur lrsquoeacutecologie des milieux serpentiniques Noumeacutea 31juillet ndash 5 aoucirct 1995rsquo Documents scientifiques et techniques III2(Centre ORSTOM de Noumeacutea Noumeacutea New Caledonia)

JanssensTKSRoeloefsDvanStraalenNM(2009)Molecularmechanismsofheavy metal tolerance and evolution in invertebrates Insect Science 163ndash18 doi101111j1744-7917200900249x

Jourdan H (1997) Are serpentine biota free from successful biologicalinvasions Southern New Caledonian ant community example InlsquoProceedings of the 2nd InternationalConference on SerpentineEcologyrsquo (Eds T Jaffreacute RD Reeves T Becquer) pp 107ndash108(ORSTOM Noumeacutea)

Karim R Van der Ent A (2014) Vegetation on ultramafic soils in KinabaluPark (Sabah Malaysia) in relation to soil chemistry and altitude InlsquoOfficial Programme Book (poster presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 42 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-poster

Kay KM Ward KL Watt LR Schemske DW (2011) Plant speciation InlsquoSerpentine the evolution and ecology of a model systemrsquo (Eds SPHarrison N Rajakaruna) pp 71ndash95 (University of California PressBerkeley CA)

Keeling SM Stewart RB Anderson CWN Robinson BH (2003) Nickel andcobalt phytoextraction by the hyperaccumulator Berkheya coddiiimplications for polymetallic phytomining and phytoremediationInternational Journal of Phytoremediation 5 235ndash244doi101080713779223

Kolaacuter F Feacuter T Štech M Traacutevniacutecek P Duškovaacute E Schoumlnswetter P Suda J(2012) Bringing together evolution on serpentine and polyploidyspatiotemporal history of the diploid-tetraploid complex of Knautiaarvensis (Dipsacaceae)PLoSONE7 doi101371journalpone0039988

Kosugi A Tamaru J Gotou K Furihata H Shimizu A Kawabe A Harada E(2015) Metal accumulation by Arabidopsis halleri subsp gemmifera at alimestone mining site Australian Journal of Botany 63 134ndash140doi101071BT14242

Kruckeberg AR (1984) lsquoCalifornia serpentines flora vegetation geologysoils and management problemsrsquo (University of California PressBerkeley CA)

Lau JA McCall AC Davies KF McKay JK Wright JW (2008) Herbivoresand edaphic factors constrain the realized niche of a native plant Ecology89 754ndash762 doi10189007-05911

Li Z Wu L Hu P Luo Y Zhang H Christie P (2014) Repeatedphytoextraction of four metal-contaminated soils using the cadmiumzinc hyperaccumulator Sedum plumbizincicola Environmental Pollution189 176ndash183 doi101016jenvpol201402034

Losfeld G Grison C Jaffreacute T Mathieu R Fogliani B LrsquoHuillier L (2014a)Leaf-age effect a key factor for reporting trace elementhyperaccumulation by plants and design applications In lsquoOfficialProgramme Book (poster presentations) 8th International Conferenceon Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD ReevesG Echevarria L LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJPollard M Lakim A van der Ent M Suleiman JB Sugau) p 23 (SabahParks Kota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Losfeld G LrsquoHuillier L Fogliani B Jaffreacute T Grison C (2014b) Miningin New Caledonia environmental stakes and restoration opportunities

Environmental Science and Pollution Research International 225592ndash5607 doi101007s11356-014-3358-x

Losfeld G Grison C Mathieu R LrsquoHuillier L Fogliani B (2014c)Phytoextraction from mine spoils insights from New Caledonia InlsquoOfficial Programme Book (oral presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 39 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Majapun R Khoo E Jumian J Jaimin AJ Tuyok M Sugau JB (2014)Dipterocarps in the eastern part of Sungai Imbak Virgin Jungle ReserveSabah In lsquoOfficial Programme Book (poster presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 36 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-poster

Malaisse F (2014) The copper-cobalt flora of Katanga and Northern Zambiarecent progress (2006ndash2014) In lsquoOfficial Programme Book (oralpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 77 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Mandal A Purakayastha TJ Patra AK (2014) Phytoextraction of arseniccontaminated soil by Chinese brake fern (Pteris vittata) effect on soilmicrobiological activities Biology and Fertility of Soils 50 1247ndash1252doi101007s00374-014-0941-8

Maroto-Valer MM Fauth DJ Kuchta ME Zhang Y Andresen JM (2005)Activation of magnesium-rich minerals as carbonation feedstockmaterials for CO2 sequestration Fuel Processing Technology 861627ndash1645 doi101016jfuproc200501017

Maycock CR Khoo E Kettle CJ Pereira JT Sugau JB Nilus R Jumian JBurslem DFRP (2012) Using high resolution ecological niche models toassess the conservation status of Dipterocarpus lamellatus andDipterocarpus ochraceus in Sabah Malaysia Journal of ForestScience 28 158ndash169 doi107747JFS2012283158

Mayer MS Soltis PS Soltis DE (1994) The evolution of the Streptanthusglandulosus complex (Cruciferae) genetic divergence and gene flow inserpentine endemics American Journal of Botany 81 1288ndash1299doi1023072445405

McAlister R Kolterman D Pollard A (2015) Nickel hyperaccumulation inpopulations of Psychotria grandis (Rubiaceae) from serpentine and non-serpentine soils of Puerto Rico Australian Journal of Botany 63 85ndash91doi101071BT14337

McCollom TM Seewald JS (2013) Serpentinites hydrogen and lifeElements 9 129ndash134 doi102113gselements92129

Medeiros ID FrydayAMRajakarunaN (2014)Additional lichen records andminerological data frommetal-contaminated sites inMaineRhodora116323ndash347 doi10311913-26

Medeiros IDRajakarunaNAlexanderEB (2015)Gabbro soilndashplant relationsin the California Floristic Province Madrono 62 75ndash87

Mesjasz-Przybyłowicz J Przybyłowicz W (2011) PIXE and metalhyperaccumulation from soil to plants and insects X-RaySpectrometry 40 181ndash185 doi101002xrs1304

Mesjasz-Przybylowicz J PrzybylowiczW (2014) Nickel hyperaccumulatorsfrom South Africa ndash present status and research challenges In In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd S StraussNRRajakarunaAJ PollardM Lakim A van der Ent M Suleiman JB Sugau) p 52 (Sabah Parks

12 Australian Journal of Botany A van der Ent et al

Kota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Mincey K Boyd RS (2014) Tolerance as an herbivore defense in thenickel hyperaccumulator Streptanthus polygaloides In lsquoOfficialProgramme Book (poster presentations) 8th International Conferenceon Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD ReevesG Echevarria L LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJPollard M Lakim A van der Ent M Suleiman JB Sugau) p 18 (SabahParks Kota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

MizunoTKirihataY (2015)Elemental compositionof plants fromserpentinesoil of Sugashima Island Japan Australian Journal of Botany 63doi101071BT14226

Mizuno T Emori K Tomioka R Harada E Takenaka C Ito SI (2014)Manganese hyperaccumulation in Chengiopanax sciadophylloides(Araliaceae) from uncontaminated soil and its correlation with calciumaccumulation In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 23 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Mohtadi A Ghaderian SM Jamali Hajiani N (2014) Geobotanical andbiogeochemical reconnaissance of the serpentine soils of Rezvanshahrin northwestern Iran In lsquoOfficial ProgrammeBook (poster presentations)8th International Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 26 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-poster

Moore MJ Mota JF Douglas NA Olvera HF Ochoterena H (2014) Theecology assembly and evolution of gypsophile floras In lsquoPlant ecologyand evolution in harsh environmentsrsquo (Eds N Rajakaruna RS Boyd THarris) pp 97ndash128 (Nova Science Publishers Hauppauge NY)

Moores EM (2011) Serpentinites and other ultramafic rocks why they areimportant for Earthrsquos history and possibly for its future In lsquoSerpentine theevolution and ecology of a model systemrsquo (Eds SP Harrison NRajakaruna) pp 3ndash28 (University of California Press Berkeley CA)

Mori T (2014) Lower greenhouse gas emissions from serpentine rock soilsthan sedimentary rock soils on undisturbed forests in Mount KinabaluBorneo In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 43 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

MoyleLLevineMStantonMWright JW(2012)Hybrid sterility over tensofmeters between ecotypes adapted to serpentine and non-serpentine soilsEvolutionary Biology 39 207ndash218 doi101007s11692-012-9180-9

Murren CJ Douglass L Gibson A Dudash M (2006) Individual andcombined effects of CaMg ratio and water availability on traitexpression in Mimulus guttatus Ecology 87 2591ndash2602doi1018900012-9658(2006)87[2591IACEOM]20CO2

Nilus R Sugau JB Chung AYC Pudente C Pereira JT Sabran S Kugan F(2014)Biodiversity richness of ultramafic forest at TawauForest ReserveSabah Malaysia In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJMBaker RDReeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 87 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Nyee-Fan L Jole SE Suleiman M (2014) Phylogenetic study of genusCleisocentron (Orchidaceae) in Sabah Malaysia In lsquoOfficialProgramme Book (poster presentations) 8th International Conferenceon Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD ReevesG Echevarria L LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJPollard M Lakim A van der Ent M Suleiman JB Sugau) p 28 (SabahParks Kota Kinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

OrsquoDell RE Rajakaruna N (2011) Intraspecific variation adaptation andevolution In lsquoSerpentine evolution and ecology of a model systemrsquo(Eds SP Harrison N Rajakaruna) pp 97ndash137 (University of CaliforniaPress Berkeley CA)

Ogar A Sobczyk L Stochmal A Stojakowska A Karlsson S Turnau K(2014)Microbial partners for phytoremediationndashHieraciumpilosella as amodel plant In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 28 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

Oline DK (2006) Phylogenetic comparisons of bacterial communitiesfrom serpentine and nonserpentine soils Applied and EnvironmentalMicrobiology 72 6965ndash6971 doi101128AEM00690-06

OstevikKLMoyersBTOwensGLRiesebergLH (2012) Parallel ecologicalspeciation in plants International Journal of Ecology 2012 939862doi1011552012939862

Palm ER Van Volkenburgh E (2014) Physiological adaptations of plants toserpentine soil In lsquoPlant ecology and evolution in harsh environmentsrsquo(Eds N Rajakaruna RS Boyd T Harris) pp 129ndash148 (Nova SciencePublishers Hauppauge NY)

Palomino M Kennedy PG Sims EL (2007) Nickel hyperaccumulation as ananti-herbivore trait considering the role of tolerance to damagePlant andSoil 293 189ndash195 doi101007s11104-007-9236-2

Pasari JR Hernaacutendez DL Zavaleta ES (2014) Interactive effects of nitrogendeposition and grazing on plant species composition in a serpentinegrassland Rangeland Ecology and Management 67 693ndash700doi102111REM-D-13-001161

PaukovAG (2009) The lichenflora of serpentine outcrops in theMiddleUralsof Russia Northeastern Naturalist 16 341ndash350doi1016560450160525

Paukov A Teptina A Morozova M (2014) Heavy metal uptake by lichenswith different secondary chemistry on ultramafic rocks In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd SStraussNRRajakarunaAJPollardMLakimAvanderEntMSuleiman JBSugau)p 61 (SabahParksKotaKinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

PavlovaDKaradjova IKrasteva I (2015)Essential and toxic element contentin Hypericum perforatum L from serpentine and non-serpentine soilsin Bulgaria Australian Journal of Botany 63 152ndash158doi101071BT14260

Pereira JT Sugau JB Nilus RDamit A Sabran S (2014) Botanical highlightsof Mount Silam Sabah In lsquoOfficial Programme Book (posterpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 35 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Pollard AJ Smith JAC (2014) Population-level variation in nickel toleranceand hyperaccumulation in Alyssum serpyllifolium from the IberianPeninsula In lsquoOfficial Programme Book (oral presentations) 8th

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 13

International Conference on Serpentine Ecology (9ndash13 June 2014)rsquo (EdsAJMBaker RDReeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 71 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Pollard AJ Reeves RD Baker AJM (2014) Facultative hyperaccumulationof heavy metals and metalloids Plant Science 217ndash218 8ndash17doi101016jplantsci201311011

Porembski S (2014) Plant diversity and conservation of granitoid inselbergsand ironstone outcrops inAfrica andMadagascar In lsquoOfficial ProgrammeBook (poster presentations) 8th International Conference on SerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarriaLLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M Lakim Avan der Ent M Suleiman JB Sugau) p 56 (Sabah Parks Kota KinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Power IM Wilson SA Dipple GM (2013) Serpentinite carbonation forCO2 sequestration Elements 9 115ndash121doi102113gselements92115

Proctor J (2003) Vegetation and soil and plant chemistry on ultramafic rocksin the tropical Far East Perspectives in Plant Ecology 6 105ndash124doi1010781433-8319-00045

Proctor J Lee YF Langley AM Munro WRC Nelson T (1988) Ecologicalstudies onGunungSilam a small ultrabasicmountain in SabahMalaysiaI Environment forest structure and floristics Journal of Ecology 76320ndash340 doi1023072260596

Przybyłowicz WJ Mesjasz-Przybyłowicz J Migula P Nakonieczny MAugustyniak M Tarnawska M Turna K Ryszka P Orłowska EZubek S Głowacka E (2005) Micro-PIXE in ecophysiology X-RaySpectrometry 34 285ndash289 doi101002xrs826

PrzybyłowiczWMesjasz-Przybyłowicz JBarnabasAVanderEntA (2014)Elemental mapping in tropical nickel hyperaccumulators from SabahMalaysia using nuclear microprobe (micro-PIXE) In lsquoOfficialProgramme Book (oral presentations) 8th International Conference onSerpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves GEchevarria LLrsquoHuillier RSBoyd SStraussNRRajakarunaAJ PollardMLakimAvanderEntMSuleiman JBSugau)p 67 (SabahParksKotaKinabalu Malaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Quimado MO Fernando ES Trinidad LC Doronila A (2015) Nickelhyperaccumulating species of Phyllanthus (Phyllanthaceae) from thePhilippines Australian Journal of Botany 63 103ndash110doi101071BT14284

Rajakaruna N (2004) The edaphic factor in the origin of plant speciesInternational Geology Review 46 471ndash478doi1027470020-6814465471

Rajakaruna N Baker AJM (2004) Serpentine a model habitat for botanicalresearch in Sri Lanka Ceylon Journal of Science 32 1ndash19

Rajakaruna N Bohm BA (2002) Serpentine and its vegetation a preliminarystudy from Sri Lanka Journal of Applied Botany 76 20ndash28

Rajakaruna N Boyd RS (2009) Soil and biota of serpentine a world viewProceedings of the sixth international conference on serpentine ecologyNortheastern Naturalist 16(Special Issue 5) 1ndash7

RajakarunaN BoydRS (2014) Serpentine soils In lsquoOxford bibliographies inecologyrsquo (Ed D Gibson) (Oxford University Press New York)

Rajakaruna N Baldwin BG Chan R Desrochers AM Bohm BA Whitton J(2003) Edaphic races and phylogenetic taxa in the Lasthenia californicacomplex (Asteraceae Heliantheae) an hypothesis of parallel evolutionMolecular Ecology 12 1675ndash1679doi101046j1365-294X200301843x

Rajakaruna N Harris TB Alexander EB (2009) Serpentine geoecology ofeastern North America a review Rhodora 111 21ndash108doi10311907-231

Reeves RD (2003) Tropical hyperaccumulators of metals and their potentialfor phytoextraction Plant and Soil 249 57ndash65doi101023A1022572517197

Reeves RD (2014) Phytomining ndash The Real Issues In lsquoOfficial ProgrammeBook (oral presentations) 8th International Conference on SerpentineEcology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G EchevarriaL LrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimA van der EntMSuleiman JB Sugau) p 22 (Sabah Parks KotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Reeves RD LaidlawW Doronila A Baker AJM Batianoff G (2015) Erratichyperaccumulation of nickel with particular reference to the Queenslandserpentine endemic Pimelea leptospermoides FMueller AustralianJournal of Botany 63 119ndash127 doi101071BT14195

Repin R (1998) Serpentine ecology in SabahMalaysia Sabah Parks Journal1 19ndash28

Roos MC Keszligler PJ Robbert Gradstein S Baas P (2004) Species diversityandendemismoffivemajorMalesian islandsdiversity-area relationshipsJournal of Biogeography 31 1893ndash1908doi101111j1365-2699200401154x

Rue M Vallance J Echevarria G Rey P Benizri E (2015) Phytoextractionof nickel and rhizosphere microbial communities under mono- ormultispecies hyperaccumulator plant cover in a serpentine soilAustralian Journal of Botany 63 92ndash102 doi101071BT14249

Sabran S Soepadmo E Leong YT Suleiman M (2014) Three Borneanendemic species of Ternstroemia (Pentaphylaceae) found in ultramafichabitats in Sabah Malaysia In lsquoOfficial Programme Book (posterpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 33 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Saibeh K Yen LV Lye Hin HC (2014) Potential of Typha angustifolia L inphytoremediation of the former Mamut copper mine Ranau SabahIn lsquoOfficial Programme Book (oral presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 30 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Selby JP Jeong AL Toll K Wright KM Lowry DB (2014) Methods anddiscoveries in the pursuit of understanding the genetic basis of adaptationto harsh environments in Mimulus In lsquoPlant ecology and evolution inharsh environmentsrsquo (Eds N Rajakaruna RS Boyd T Harris)pp 243ndash266 (Nova Science Publishers Hauppauge NY)

SeneviratneMSeneviratneGMadawalaHMSP IqbalMCMRajakarunaNVithanageM(2015)Apreliminary studyof the roleofbacterialndashfungal co-inoculation on heavy metal phytotoxicity in serpentine soil AustralianJournal of Botany 63 doi101071BT14270

Siebert S (2014) An overview of serpentine ecology research in southernAfrica and the way forward In lsquoOfficial Programme Book (oralpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 49 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Smith DC Siebert SJ Claassens S Swemmer T (2014) Relationship betweensoil microbial properties plant cover and biomass on ultramafic tailingsfacilities In lsquoOfficial Programme Book (oral presentations) 8thInternational Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S

14 Australian Journal of Botany A van der Ent et al

Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 32 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

SouthworthD TackaberryLEMassicoteHB (2014)Mycorrhizal ecology onserpentine soils Plant Ecology amp Diversity 7 445ndash455doi101080175508742013848950

Spasojevic MJ Damschen EI Harrison S (2014) Patterns of seed dispersalsyndromes on serpentine soils examining the roles of habitat patchinesssoil infertility andcorrelated functional traitsPlantEcologyampDiversity7401ndash410 doi101080175508742012678506

Sumail S Van der Ent A (2014) The ultramafic Nepenthes of Kinabalu ParkIn lsquoOfficial Programme Book (poster presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJM BakerRD Reeves G Echevarria L LrsquoHuillier RS Boyd S Strauss NRRajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 43 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-poster

TeptinaAPaukovA (2014)Nickel andothermetals uptake and accumulationby species of Brassicaceae from the ultramafic rocks in the Urals InlsquoOfficial Programme Book (oral presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 62 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

Tjoa A Barus H (2014) Heavy metals and other elemental concentrations inAsteraceae grown in top and overburden soils of serpentine fromSorowako Indonesia In lsquoOfficial Programme Book (posterpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 27 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

Turner TL Von Wettberg EJ Nuzhdin SV (2008) Genomic analysis ofdifferentiation between soil types reveals candidate genes for localadaptation in Arabidopsis lyrata PLoS ONE 3 e3183doi101371journalpone0003183

Turner TL Bourne EC Von Wettberg EJ Hu TT Nuzhdin SV (2010)Population resequencing reveals local adaptation of Arabidopsis lyratato serpentine soils Nature Genetics 42 260ndash263 doi101038ng515

USFishandWildlifeService (1998) lsquoRecoveryplan for serpentine soil speciesof the San Francisco Bay Arearsquo (US Fish andWildlife Service PortlandOR)

van der Ent A Reeves RD (2015) Foliar metal accumulation in plants fromcopper-rich ultramafic outcrops case studies from Malaysia and BrazilPlant and Soil 389 401ndash418 doi101007s11104-015-2385-9

van der Ent A Baker AJM Reeves RD Pollard AJ Schat H (2013a)Hyperaccumulators of metal and metalloid trace elements facts andfiction Plant and Soil 362 319ndash334 doi101007s11104-012-1287-3

van der Ent A Baker AJM Van Balgooy MMJ Tjoa A (2013b) Ultramaficnickel laterites in Indonesia (Sulawesi Halmahera) mining nickelhyperaccumulators and opportunities for phytomining Journal ofGeochemicalExploration128 72ndash79 doi101016jgexplo201301009

van der Ent A Repin R Sugau J Wong KM (2014a) lsquoThe ultramafic flora ofSabah an introduction to the plant diversity on ultramafic soilsrsquo (NaturalHistory Publications (Borneo) Kota Kinabalu Malaysia)

van der Ent A Erskine P Sumail S (2014b) Habitat and plantndashsoilrelationships of nickel hyperaccumulators from ultramafic soils inSabah (Malaysia) In lsquoOfficial Programme Book (oral presentations)8th International Conference on Serpentine Ecology (9ndash13 June2014)rsquo (Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RSBoyd S Strauss NR Rajakaruna AJ Pollard M Lakim A van der Ent M

Suleiman JB Sugau) p 55 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

van der Ent A Wong KM Sugau J Repin R (2015) Plant diversity ofultramafic outcrops in Sabah (Malaysia) with a focus on KinabaluPark Australian Journal of Botany 63 doi101071BT14214

Varela RP Garcia GAA Ambal AM Gonzales NP (2014) Ecobeltestablishment employing agroforestry concepts in the progressiverehabilitation of nickel mining areas towards ecorestoration InlsquoOfficial Programme Book (oral presentations) 8th InternationalConference on Serpentine Ecology (9ndash13 June 2014)rsquo (Eds AJMBaker RD Reeves G Echevarria L LrsquoHuillier RS Boyd S StraussNR Rajakaruna AJ Pollard M Lakim A van der Ent M Suleiman JBSugau) p 75 (Sabah Parks Kota Kinabalu Malaysia) Available fromhttpwwwicse2014comicse2014abstract-booklet-oral

VenterALevanetsASiebert SRajakarunaN(2015)Apreliminary surveyofthe diversity of soil algae and cyanoprokaryotes on mafic and ultramaficsubstrates in South Africa Australian Journal of Botany 63doi101071BT14207

Veter NM DeSantis LRG Yann LT Donohue SL Haupt RJ Corapi SEFathel SL Gootee EL Loffredo LF Romer JL Velkovsky SM (2013) IsRapoportrsquos rule a recent phenomenon A deep time perspective onpotential causal mechanisms Biology Lettersdoi101098rsbl20130398

Viehweger K (2014) How plants cope with heavy metals Botanical Studies(Taipei Taiwan) 55 35 doi1011861999-3110-55-35

Von Wettberg EJ Wright JW (2011) Genomic approaches to understandingadaptation In lsquoSerpentine the evolution and ecology of a model systemrsquo(Eds SP Harrison N Rajakaruna) pp 139ndash153 (University of CaliforniaPress Berkeley CA)

VonWettberg EJ Ray-Mukherjee J DrsquoAdeskyN NesbethD Sistla S (2014)The evolutionary ecology and genetics of stress resistance syndrome(SRS) traits revisiting Chapin Autumn and Pugnaire (1993) In lsquoPlantecology and evolution in harsh environmentsrsquo (Eds N Rajakaruna RSBoydTHarris) pp 201ndash226 (NovaSciencePublishersHauppaugeNY)

Whiting SN Reeves RD Richards D Johnson MS Cooke JA Malaisse FPaton A Smith JAC Angle JS Chaney RL Ginocchio R Jaffreacute T JohnsRMcIntyre T PurvisOW Salt DE Schat H Zhao FJ BakerAJM (2004)Research priorities for conservation ofmetallophyte biodiversity and theirpotential for restoration and site remediation Restoration Ecology 12106ndash116 doi101111j1061-2971200400367x

Whitman M Russo SE (2014) Does Rapoportrsquos rule apply to the ultramaficflora of Sabah Borneo In lsquoOfficial Programme Book (posterpresentations) 8th International Conference on Serpentine Ecology(9ndash13 June 2014)rsquo (Eds AJM Baker RD Reeves G Echevarria LLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M LakimAvanderEntMSuleiman JBSugau)p 54 (SabahParksKotaKinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-poster

WijethungaWMKTWijesekara KB Weerasooriya R Rajakaruna N (2014)Chromium phytoextraction potential of Brassica juncea (L) Czern(Indian mustard) genotypes from Sri Lanka In lsquoOfficial ProgrammeBook (oral presentations) 8th International Conference on SerpentineEcology (9ndash13June2014)rsquo (EdsAJMBakerRDReevesGEchevarriaLLrsquoHuillier RS Boyd S Strauss NR Rajakaruna AJ Pollard M Lakim Avan der Ent M Suleiman JB Sugau) p 17 (Sabah Parks Kota KinabaluMalaysia) Available from httpwwwicse2014comicse2014abstract-booklet-oral

Wong KM (1992) Sabahrsquos plant life a new look at a priceless wonder InlsquoThe environment the future is in our handsrsquo (Ed Anon) pp 26ndash30(Intan Junior Chamber Kota Kinabalu Malaysia)

Wright JW Stanton ML (2011) Local adaptation in heterogeneouslandscapes reciprocal transplant experiments and beyond InlsquoSerpentine the evolution and ecology of a model systemrsquo (Eds SP

Global research on ultramafic (serpentine) ecosystems Australian Journal of Botany 15

Harrison N Rajakaruna) pp 155ndash179 (University of California PressBerkeley CA)

Wu CA Lowry DB Cooley AM Wright KM Lee YW Willis JH (2008)Mimulus is an emergingmodel system for the integrationof ecological andgenomic studies Heredity 100 220ndash230 doi101038sjhdy6801018

Wulff AS Hollingsworth PMAhrends A Jaffreacute T Veillon JM LrsquoHuillier LFogliani B (2013) Conservation priorities in a biodiversity hotspotanalysis of narrow endemic plant species in New Caledonia PLoSONE 8(9) e73371 doi101371journalpone0073371

Yang H Xu Z Fan M Gupta R Slimane RB Bland AE Wright I (2008)Progress in carbon dioxide sequestration and capture a review Journal ofEnvironmental Sciences 20 14ndash27doi101016S1001-0742(08)60002-9

Yost JM Barry T Kay KM Rajakaruna N (2012) Edaphic adaptationmaintains the coexistence of two cryptic species on serpentine soilAmerican Journal of Botany 99 890ndash897 doi103732ajb1100521

ZhangX Laubie B Houzelot V Plasari E Barbaroux RMercier G Blais JFBani A Morel J-L Echevarria G Simonnot M-O (2014) New trends fornickel phytomining In lsquoOfficial Programme Book (oral presentations)8th International Conference on Serpentine Ecology (9ndash13 June 2014)rsquo(Eds AJM Baker RD Reeves G Echevarria L LrsquoHuillier RS Boyd SStrauss NR Rajakaruna AJ Pollard M Lakim A van der Ent MSuleiman JB Sugau) p 27 (Sabah Parks Kota Kinabalu Malaysia)Available from httpwwwicse2014comicse2014abstract-booklet-oral

16 Australian Journal of Botany A van der Ent et al

wwwpublishcsiroaujournalsajb