Papers and Proceedings of the Royal Society of Tasmania, Volume 143(2), 2009 87
AN ESTIMATE OF CHANGE IN A SUB-POPULATION OF LOMATIA TASMAN/CA (PROTEACEAE) BETWEEN 1993 AND 2008
by Jayne Balmer
(with seven text-figures and one table)
Balmer, J. 2009 (11 :xii): An estimate of change in a sub-population of Lomatia tasmanica (Proteaceae) between 1993 and 2008. Papers and Proceedings of the Royal Society a/Tasmania 143(2): 87-94. https://doi.org/10.26749/rstpp.143.2.87 ISSN 0080-4703. Biodiversity Conservation Branch, Department of Primary Industries, Parks, Water and Environment, GPO Box 44, Hobart, Tasmania 7001, Australia. Email: Jayne.Balmer@ dpiwe.tas.gov.au
A decline in ramet (plant stem) numbers was observed within a sub-population of Lomatia tasmanica W.M. Curtis between 1993 and 2008. The mortality was mainly among the small ramets. This resulted in a shift from a population dominated in density by recent suckers and young ramets ro one dominated by more mature ramets. lhere was possibly an increased survival rate among ramets cut forgenetic sampling in 1993 compared with those that were not pruned. It is suspected that a combination of reduced rainfall and declining light availability has resulted in an increased mortality among young ramets. Although this species is entirely reserved within the Tasmanian Wilderness World Heritage Area it is still vulnerable to random events and climate change and its status as critically endangered remains appropriate. Key Words: threatened plant monitoring, pruning impact, climatechange, Southwest National Park, Tasmania, Lomatia tasmanica.
et al. 2004). Despite the success and advantages of asexual INTRODUCTION reproduction (Cook 1985) L. tasmanicais one of only a few vascular plant taxa that are known to reproduce entirely by Lomatia tasmanica WM. Curtis (King's Lomatia) is a asexual means. A few other examples also occur within the member of the plant family Proteaceae Juss. and is listed family Proteaceae including the endangered clonal shrub as critically endangered under the Australian Government's Hakea pulvinifera L.A.S. Johnson (Smith 2004). As one of Environmental Protection and BiodiversityConservation Act just a few taxa exhibiting this trait, L. tasmanica provides 1999. The plant reproduces vegetatively by sending up new a useful example to test the evolutionary hypothesis that plant stems, defined here as ramets, from underground followingthe extinction of their sexual ancestors, clones will rhizomes (Brown & Gray 1985). In December 1993 surveys inevitably become extinct without taxonomic descendants by ]. Balmer and others mapped ail ramets within one (Grant 1981). transect region (Lynch & Balmer 2004) and sampled leaves This region of southwest Tasmania has experienced a and population densities fromacross the known geographic substantial and linear decline in rainfall over the past range of the species to determine the genetic variability of decade (Bureau of Meteorology data for Maatsuyker Island, the population and estimate the total population of the Kirkpatrick et al. 2002), with some areas recently receiving stand. The genetic study revealed that the population was less than the 50 mm summer monthly rainfall considered composed of a single genetic clone (Lynch et al. 1998). necessary to maintain rainforest (Jackson 1968). This raises Chromosome counts led to the conclusion that the species the question of how climate change might impact on plant is a sterile triploid relying entirely on vegetative regeneration dynamics within rainforest - especially small relict patches. for its survival (Lynch et al. 1998, Threatened Species This further increases the scientific and conservation Section 2005). A fossilrecord of the species fromMelaleuca management interest in monitoring L. tasmanica, one of ° ° (�43 25'5 146 l0'E) in southwest Tasmania provides the few plants to have been intensely surveyed in the region evidence that this clonal plant may have been in existence prior to the reduction in annual and summer rainfall. for 43 600 years (Jordan et al. 1991, Lynch et al. 1998). Another reason formonitoring has been the concern that Data from surveys conducted between 1993 and 1996 taking cuttings from I. tasmanica plants may cause shock provided a population estimate for the plant clone of 600 or allow disease to invade plants, leading to the death of ramets (stems) (Lynch & Balmer 2004). The total extent of sampled plants in the wild. Potential threats to the species the species is now thought to be about 1.2 km within the are posed by naturalists and gardeners illegally taking stem Southwest National Park in Tasmania (Lynch et al. 1998, samples to propagate the species for horticulture; such Threatened Species Unit 2000, Lynch & Balmer 2004). The visits also increase the potential to introduce disease into habitat for the species ranges from wet scrub to rainforest the population. with most of its extent occurring within riparian areas of In response to the threatened species recovery plan mixed forest (Brown & Gray 1985, Lynch& Balmer 2004). recommendations and concerns about both the impact of The recovery plan for I. tasmanica recommends that recent rainfall declines and the taking of cuttings from the population monitoring be undertaken "to find more out wild population the area surveyed in 1993 was resurveyed about this unique species" (Threatened Species Section in March 2008. This paper explores the relationships 2006, p. 15). Monitoring of threatened species is essential observed between the 1993 and 2008 survey data and tests to determine if populations are contracting and require the following two null hypotheses: (1) the population size management intervention in order to avert further decline and demography of the L. tasmanica stand has not changed and extinction. I. tasmanica has WorldHeritage significance between 1993 and 2008; and (2) the health of L. tasmanica as an example of the longest-lived plant clone known in the ramets from which cuttings were taken is no different from world and has had intense international publicity (Balmer plants from which stem cuttings were nor taken. 88 Balmer
METHODS
1993 (unpublished The Lynch & Balmer 2004) u~ srem length and distance to the creek were significantly c•a•.uua A 75 rn transect line was run out using a tape measure in between the 1993 that survived to 2008 compared straight line sections so rhat it more or less followed the creek with those that 2008. -lhe Chi--squared test was line. 1hc changing direction of the transect was recorded in was independent of degrees (magnetic) using a Sunto compass and the distance stem diameter and the of stem cutting. in metres to each bend was noted. All living ramets recorded in each of the two time All ramets of L tasmaniw observed within 10 m either periods within the transect area were used to rhe side of the transect line (an area of ··1.5 ha) were recorded significance of structural differences between together with: the two tirne 1ne significance of the difference in • rhe distance away hom the creek in an median values of stem diameter and of the 2008 and eastern or western direction 1993 population was tested the W the distance up from the beginning of the transect test. Linear and (~northwards) was used to test the relative importance • an estimate of height or ramet the nearest 500 the survival of mm) 'TI1e student t--test was used to compare the difference in • the stem diameter in millimetres measured at the base the mean climatic data for the 14 preceding the 1993 using a diameter tape and 2008 monitoring visits and long--term weather data • the number of upright ramets arising from a basal rhizome for Maatsuyker Island nearest weather station). All tests • notes on the plant health were performed the software f""'"'"a,;", ouu~.ru1mLu • a note recording if a cutting was taken for Windows (Manugistics 2000). • a ramet identification number. If a main stem (ramet) was obvious when leaning or horizontal) it was recorded and mapped at its point of RESULTS origin from the ground. However, where a basal horizontal stem was present very dose to the ground with more than The total number of living plant stems of L. tasmanica one main leader arising from it and behaving as individual observed in the area changed from 103 in 1993 to 84 trees then each of these main leaders was treated as an in 2008, a decline 18%. Eight ramets observed in 2008 individual ramet for the purposes of the census. were small ( <3 em diameter or <600 mm high) and to Flagging tape was tied to each L. tasmanica ramet found have regenerated since 1993 of the 2008 population). to make it clear that had been recorded. All ramets Fourteen unmarked ramets observed in 2008 could not be from which cuttings were taken and an equivalent number matched to the 1993 mapped ramet locations and were of control ramets were tagged with stainless steel measuring estimated to be larger than plants arising since 1993. Tnese labels loosely hung around the branch with the wire enclosed unmatched ramets were assumed to have been overlooked in tubing to protect the plant from abrasion or in 1993. A total of 41 plants of the 103 live plants recorded damage from the tag. in 199 3 were either dead or not located and were presumed to have died since 1993 (table l). Excluding the 2008 2008 census unmatched mature plants from the population statistics the decline in plants recorded in 1993 was around 32% (table In March 2008 all ramets at the site were mapped following If they are added to the original population numbers the the method used in 1993 and an attempt made to match decline was 28%. rarnets with the original ones located in 1993. The ramets The survival rate was among plants that stiil bore numbered metal tags or labelled flagging tape three of the 16 were matched precisely with the 1993 survey data. Due to pi' ants fron1 the transect area vvere found dead the imprecision of the mapping in 1993 and time constraints in 2008 or not relocated. The Chi square test with Yates for the remapping in 2008 it was not possible to p1<:c."'"'Y correction factor was only at the 90% level match all ramets between the two sampling (p
\
.~ Dead in 2008 ':It Dead in i 993
A. .. ,~ .~i A "" J.
* AI& A " *
.i.
*
*
FIG. I - Map of ramets in 1993 within W m the FIG. 2- of ramets zn 2008 within 10 m the transect-line. lhe inset map of Tasmania shows the location transect-line. of the transect dot) and the location of the Maatsuyker Island climate station (star).
TABLE 1 Tally of plants recorded in 1993 and 2008 ------~~--"'--- --·-----··------Population parameters Tally of live ramets of dead ramets ------·------Total no. of ramets observed in 1993 [P 1] 103 5
Total no. of ramets observed in 2008 84 11
Live 2008 1 [L] and dead 2008 [Dl] 62 11 Unmatched plants regenerated since 1993 [R] 14 0 Unmatched plants (possibly overlooked in 1993) 8 0
Ramets not relocated in 2008 presumed to have died2 [D2] 30
Regeneration rate 1993-2008 [R/live live P1] 3 8%
Mortality rate 1993--2008 [D 1 < D2 /live P1 P 40% Population change 1993-2008 [1-{ (L-+:.£::-_12_1-Q~)/Iive Pl}] ·1 -32%
1 Two live and one dead ramet were marked but did not match locations of the plants mapped in 1993. 2 lhe marked ramets located in areas not matching the original mapped locations were subtracted from the tally of missing plants. 3 Calculations exclude unmarked mature ramets observed in 2008 not observed in 1993. 90 Balmer
1 'm) . 1. 30 20 ,_{'() '10 0 10 20 30 0 2 3 4 5 5
FIG. 3 --·-· the 1993 rarnets that F!G 4 --- the 1993 that died by 2008.
30 20 10 0 10 20 30 40 0 2 3 4 5 0 20 40 60 80 100 '! 1
FIG. 5 - 1-firfnrn•arn HG. 6--- 1-!ictnm·.-um diameters between 1993 and 2008. between 1993 and 2008.
F!G.7 station 1979to2007. An estimate ofchange in a sub-population ofLomatia tasmanica (Proteaceae) between 1993 and 2008 91
p<0.05) but stem diameter did not provide any additional 600 ramets to betl;yeen 400 and 500 ramets. In addition explanation of the variation (p>0.05). an ex-sim collection of plants is maintained by the Royal The median diameter and height of ramets was significantly Tasmanian Botanical Gardens (RTBG) with only a v~ry different between the two time periods (p
Possible explanations for population decline of variance suggests that it was the with the greatest leaf area or leaves higher above the ground that were more Given that the taking of stem cuttings appears unrelated likely to survive. 1hese plants are likely to intercept a greater to the observed pattern of mortality, other exphnadons are amoum of light. required for the population decline observed within the Ihe vigour of plants in the wild is poor to plants transect area. 'TI1e following hypotheses are explored here: grown in gardens and nurseries where light is more fieely • introduction of a plant pathogen such as available and competition is reduced suggesting that light cinnamomi may be a limiting factor in the wild. Nevertheless the species • competition for light is very sun-sensitive and shade (Mark Fountain and e moisture stress Natalie Tapson pers. comm. 2008)" A study of ramet • disturbance from trampling during the 1993 survey. response to light variation in a species of bamboo showed that low light conditions inhibited the production of ramets, Pathogen and resulted in ramets with longer internode bur no mortality was recorded et al. Pathogenicity tests have shown thatl. tasmanicais susceptible to 1? cinnamomi (Rudman 2000). 1his plant pathogen is Moisture stress known to occur within the buttongrass moorland vegetation adjoining the forests in which L. tasmanica occurs but no The average rainfall fiJr the period tests of the soil within the stand or the creek water within up to 2008 was c.l 000 mrn th;n the the stand have been undertaken to determine if the pathogen preceding 14-year period it fluctuated between 1200 has invaded the stand itself (Tim Rudman, Department of and 1800 mm (fig. 7). This has shifted the rainfall doser to Prim.ary Industries, Parks, Water and Environ~ent Files, 8 the lower limit for the maintenance of rainforest vegetation March 2001). of about 1000 rnm average annual per year and 50 mm It is unlikely that the recent decline in ramets could be average summer monthly rainfall 1968). 1he higher attributable to this pathogen since studies of the summer than average rainfall experienced in the years leading up ro soil temperatures in closed forest habitat similar to that 199 3 may have given rise to an increased production of along the transect have shown that these soils are too cool rarnets that have subsequently died. Kirkpatrick et al. (2002) to support the growth and reproduction of P cinnamomi suggested that a decline in graminoid and herb cover in the (Podger & Brown 1989, Rudman 2000). Analysis of recent alpine vegetation on the Southern Ranges between 1990 average monthly maximum temperature data for the region and 2000 may have been related to the significant linear showed no significant changes in air temperatures, which decline in rainfall between 1980 and 2000. No other is an indication that summer soil temperatures have not in southwest Iasmania has looked at vegetation responses to increased substantially in recent times although the recent climatic ~'"'H";'-'· of averaged monthly data may conceal more subtle changes Reduced moisture availability is known to reduce the in climate. Nevertheless, no evidence for 1? cinnamomi, production of proteoid roots in some members of the such as ill health or death of other susceptible species, was Proteaceae (Lamont In general, with proteoid observed within the rainforest during the various surveys roots are able to extract phosphorous from nutrient-deficient and visits to the stand. soils more effectively than other plants, even those with All visitors to the stand have been asked to follow strict mychorrhizal associations (Pearson & Rengel 1 If hygiene procedures before entering the forest and visitation is L tasmanica has such roots it is that if moisture levels discouraged. Ihis protocol is applied widely in conservation are reduced then the plant's to absorb phosphorous reserves where values are at risk from the of this will decline, its ability in this low- pathogen (Cahill eta/. 2008). Here it aims to reduce the nutrient environment. chance of introducing long-lived P cinnamomi into habitat occupied by L. tasmanica. A disturbance opens Spatial distribution plant the canopy, such as fire, could enable soil temperatures to increase sufficiently for P cinnamomi to become active leadinr: 1he spatial pattern of the dead plants relative to the living to the death of ~usceptible species such as L tasmanic~ population was not tesred for its deviation from as they regenerate after the fire (Podger & Brown 1989, random. The disturbance from survey work is Rudman 2000). been distributed across the or concentrated in the of the creek-line Light availability transect-line was located. the deaths appear to be d ustered in patches rather than being evenly dispersed Since the last fire at the site in 1934, the rainforest or concentrated along the so there is no apparent have become increasingly dominant in the understorey and relationship between the site survey conducted in 1993 and it is likely that this has reduced light penetration to the the of deaths observed. ground layer. Accumulating iitter and branch wood, as well as with proteoid roots, such those in the genus competition from other ground layer species, may be factors Lomatia and many other members of the Proteaceae (Purnell contributing to the higher mortality of small ramets and the 1 be sensitive to root disturbance reduction in the establishment of new suckers. Data were the disturbance of potting up not collected on relative light plant competition them is associated with an or understorey litter, branch wood or tree falls so that this increased mortaiity (Lorraine Perrins cornrn. July hypothesis was unable to be tested. Nevertheless, the greater 2008 and A recent death the most mature importance of height for survival and the relative lack of plant in the RTBG collection foliowed the removal of importance of stern diameter in the multi-.factor uPonnAPrc from the soil surface. Tests did not reveal the cailSe An estimate 93 of the death but the symptoms of sudden blackening of ail national park in the Tasmanian Wilderness World Heritage the leaves suggesred that the plant may have been shocked Area does not offer any protection this threat. Other by the light root disturbance the release of toxic species with limited dispersal and sensitivity to phenolic compounds which led to its death. drought are also likely to be at risk from declines in local Given the apparent this species to root regional rainfalL While climate modelling for the Tasmanian disturbance some impact from during the survey and Australian region has not predicted any significant of 1993 cannot be dismissed. changes in the climate of Tasmania it is based on limited data and hence has a low reliability for this Clonal plant responses to resource region et al. 2005, CSIRO & the Australian Bureau of Meteorology 2007). Each ramet of L tasmanica is part of a larger colonial plant. Given the threats to the wild population of L. tasmanica, 1he duration of the connections beCV'reen L tasmanica the maintenance of an ex-situ population of the species is ramets is not known but studies of showed that important Cll1reatencd Section 2006). Although root connections between donal trees may for up a clone, the low rate of survival of plants in horticulture to 50 while others & Ashmun suggests that rhe ex-situ collection should be no less than Clonal plants can, 50 plants. The establishment of lived material (Cook 1985, Pitelka & Ashmun 'TI1ey are atso able grafting onto root stock also be a to "forage" altering their growth in response to resource precaution. 'The development of a cheap, mass propagation availability, despite this their success may be dependent technique for the species would enhance the long-term on the spatial arrangement of suitable habitat et al. survival chances of the species 1996). Observations and ecological about dynamics • the total number of extant plants in horticulture of donal populations suggest that the parent plants e reducing the need to visit the wild population in order expend energy to support their daughters and that to see the species to a higher rate of establishmem by clones compared with • reducing the illegal taking of cuttings from the wild seedlings (Thomas & Dale 1975, Cook 1985). Populations • enabling the efficient propagation of the species for are fairly stable over long periods of time with increases replanting in the event of a wildfire or any other in appearance of new plants tending to be associated with catastrophic event destroying the wild population. increased rates of mortality which are typically seasonal A first attempt to develop a tissue culturing technique (Noble et al. 1979, Fletcher & Shaver 1983, Cook for the species was begun at the University ofTasmania in More recent theoretical models suggest that habitat patchiness 1994 (Cambecedes 1 This research is continuing as a and variabiiity is likely w determine whether the strategy collaborative project between the University of Tasmanian of maintaining integration between clonal and the RTBG (Anthony Katoulis and Lorraine Perrins advantageous to the the pers. comm. April and the project's continuation is progeny from parent plants (Kun & recommended. et al. Since the degree to which L tasmanica Further ecological studies ofL tasmanica are also desirable or remains integrated is unlmown, the implications if there is to be any chance of understanding and conserving on its capacity to respond successfully to changes in habitat this unique species in the wild. Priority research questions remain unclear. include: • Does L tasmanica have proteoid roots and what ramifications does this have for horticulture? CONCLUSIONS • How sensitive to trampling impacts is the plant and what measures can be taken to minimise impacts of visitation 'The results of this to conservation to the wild population? management of longitudinal studies to monitor How long do ramets maintain connections with each the population of threatened and the risks other? "This has ramifications on stand maintenance too much reliance on the through resource sharing and partitioning. highlight that threatened are • Is light or other competition a limiting factor for and outside the reserve system. Species restricted to the regeneration of the species in the wild and if so could it reserve system may be vulnerable to benign neglect by land be alleviated active canopy thinning or removing of managers struggling to find sufficient funds to manage the competing dead branchwood and other litter? reserve system .. • What prompts the formation of new ramets, and what that the decline in numbers of ramets is their growth rate? nh,pr·vpd in the transect area is part of a natural rP<:ncm<:e @ Are current summer soil temperatures sufficiently low to to climatic variation and/or A study protect the species from being impacted by size class structure of L tasmanica across its range is needed cinmzmomi should it invade? to determine if the decline observed in this sub-popLllation • Are spores of P cinnamomi in the soil within the is of a rnore trend. L tasmanica stand? there has been a decline in younger the survival of the mature suggests that the sub-population is not under immediate risk oflocal extinction. ACKNOWLEDGEMENTS is still albeit at a lower rate than previous If declining rainfall is Lynch and Karen Johnson took part in the field survey then further 1993 and 2008 respectively. Colin Reid provided GIS the rPO"PnPn assistance with the original mapping of the 1993 data. This survival of L tasmanica in. the project was funded the Australian Government through 94 jayne Balmer the Tasmanian Wilderness \Vorld Heritage Area program. an ancient done. ofBotany 46: 2 5-33. The climatic data were supplied the Australian Bureau of Mcintosh, P., Pook, M. & McGregor, J. 2005: Study of future Meteorology. 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