Distribution, habitat, and relation to climatic factors

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Distribution, habitat, and relation to climatic factors of the lichen genus Ramalina in New Zealand a

a

Peter Bannister , Jennifer M. Bannister & Daniel J. Blanchon

b

a

Department of Botany, University of Otago, P.O. Box 56, Dunedin, New Zealand b

Resource Management Research Group, School of Landscape and Plant Science UNITEC, Private Bag 92025, Auckland, New Zealand Version of record first published: 17 Mar 2010.

To cite this article: Peter Bannister , Jennifer M. Bannister & Daniel J. Blanchon (2004): Distribution, habitat, and relation to climatic factors of the lichen genus Ramalina in New Zealand, New Zealand Journal of Botany, 42:1, 121-138 To link to this article: http://dx.doi.org/10.1080/0028825X.2004.9512894

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New Zealand Journal of Botany, 2004, Vol. 42: 121-138 0028-825X/04/4201-0121 © The Royal Society of New Zealand 2004

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Distribution, habitat, and relation to climatic factors of the lichen genus Ramalina in New Zealand

PETER BANNISTER JENNIFER M. BANNISTER Department of Botany University of Otago P.O. Box 56 Dunedin, New Zealand DANIEL J. BLANCHON Resource Management Research Group School of Landscape and Plant Science UNITEC Private Bag 92025 Auckland, New Zealand Abstract We present maps, based on herbarium records extensively supplemented by our own records, showing the distribution of the 14 species of Ramalina found in New Zealand (excluding the Kermadec, Chatham, and sub-antarctic islands). Species diversity is highest in two areas, eastern Northland (seven species) and coastal Otago (eight species). Only four species (R. canariensis, R. celastri, R. glaucescens, and R. unilateralis) are common to both main islands. Distribution is related to climate and altitude along a latitudinal gradient on the eastern side of New Zealand, from cooler, drier sites and higher altitudes in the south to warmer, moister sites and lower altitudes in the north. Species of Ramalina tend to be absent from sites with various combinations of high rainfall, high altitude, high water deficits, and extremes of temperature. Species reproducing asexually and sexually show similar relationships of frequency to spread, but with sexually reproducing species showing greater frequency at a given degree of spread. Keywords Ramalina; Ramalina arabum; Ramalina australiensis; Ramalina canariensis; Ramalina celastri; Ramalina erumpens; Ramalina B02091; Online publication date 30 March 2004 Received 24 December 2002; accepted 23 June 2003

fimbriata; Ramalina exiguella; Ramalina geniculata; Ramalina glaucescens; Ramalina inflexa; Ramalina meridionalis; Ramalina pacifica; Ramalina peruviana; Ramalina riparia; Ramalina unilateralis; distribution; habitat; climate; New Zealand; lichens

INTRODUCTION Ramalina Ach. is a cosmopolitan genus of fruticose lichens found in locations from beyond the Arctic Circle to the Antarctic Peninsula and encompassing a range of climates from polar to tropical. Kirk et al. (2001) listed c. 200 species of Ramalina world-wide. Fourteen species of Ramalina occur in New Zealand (Blanchon et al. 1996). Species of Ramalina occurring on the outlying islands of the New Zealand geographic area are not included in this paper but are documented by Bannister & Blanchon (2003). R. arabum (Dill. ex. Ach.) Meyen & Flotow is no longer considered to occur in New Zealand and specimens described under this name constitute a new species, R. meridionalis D.Blanchon & J.Bannister (Blanchon & Bannister 2002; see also Müller Argoviensis 1894; Hellborn 1896; Zahlbruckner 1941; Martin 1966, 1968; Galloway 1985; Blanchon et al. 1996). The New Zealand species of Ramalina can be divided into three groups: those most prevalent in the warmer temperate regions of northern New Zealand (latitude 44°S), and those occurring in both North and South Islands. The northern group comprises R. meridionalis, R. australiensis Nyl., R. pacifica Asahina, R. peruviana Ach., R. exiguella Stirt., and R. geniculata Hook.f. & Taylor. The southern group consists ofR. erumpens D.Blanchon, J.Braggins & A.Stewart, R. fimbriata Krog & Swinscow, R. inflexa D.Blanchon, J.Braggins & A.Stewart, andR. riparia D.Blanchon, J.Braggins & A.Stewart. Four species occur in both North Island and South Island: R. canariensis J.Steiner andR. celastri (Spreng.) Krog & Swinscow are more frequent in North Island while


122 R. glaucescens Kremp. andR. unilateralis F.Wilson are more frequent in South Island. Six of the 14 species (R. celastri, R. exiguella, R.glaucescens, R. inflexa, R. geniculata, and R. exiguella) are consistently apothecial. R. meridionalis andR. australiensis are largely vegetative and probably spread via thallus fragmentation (Bannister & Blanchon 2003), but occasionally produce apothecia. The remaining species produce soredia; but apothecia occur occasionally on R. peruviana and rarely on R. unilateralis and R. fimbriata, but have not been observed on R. canariensis, R. erumpens, or R. pacifica in New Zealand. Lichens that spread by asexual means are generally considered to be more frequent and widespread than those that produce only sexual spores (e.g., Bowler & Rundel 1975; Louwhoff 2001). The first maps based on herbarium records (Blanchon et al. 1996) have been expanded by our own records from the field and additional records from herbarium specimens not included in earlier maps. Since September 1997, we have visited localities all over New Zealand targeting under-recorded areas and species. Our main objective was to produce more complete maps of the distribution of the New Zealand species of Ramalina and use these to relate their distribution to climate using the climate response surfaces developed by Leathwick & Stephens (1998). A secondary objective is to use the information on frequency and location to examine whether those species reproducing asexually are more frequent and more widespread than those producing sexual spores.

MATERIALS AND METHODS Blanchon et al. (1996) produced maps that relied largely on herbarium and other records collated by Blanchon (1994). Examination of these maps showed that there were areas of the country where species of Ramalina were either absent or unrecorded. Ongoing investigations began in September 1997 in the Dunedin area and Otago and were followed by specific visits to more distant areas in subsequent years. These were Stewart Island and western Fiordland, Jan 1998; Bay of Islands, Jun 1998; Taranaki-Manawatu, Nov 1998; Marlborough-Nelson and West Coast, Dec 1998; Southland, Apr 1999; Coromandel, Aug 1999; Wellington to East Cape, Nov-Dec 1999; Te Anau, Apr 2000; western and central Northland, Jun-Jul 2000; central North Island, Dec 2000; Southland,

New Zealand Journal of Botany, 2004, Vol. 42 Apr 2001; Banks Peninsula and Mid Canterbury, July 2001; North Canterbury, Aug 2001; Marlborough, Apr 2002; Great Barrier Island, Dec 2002. Additional records were obtained by examining specimens from herbaria at Auckland (AK), Wellington (WELT), Lincoln (CHR), Canterbury University (CANU), recent additions at Otago (OTA), and the private collection of P. N. Johnson. Additional specimens collected by individuals (see Acknowledgments) were verified and included in the mapping. In total, 1535 separate sites were mapped, 609 from herbarium records and 926 from our own records (Fig. 1). Grid references (New Zealand Map Grid, NZMG) and altitude were determined for each site where our records were made, and from herbarium records that gave specific map references and altitudes. Landcare Research (Hamilton, New Zealand) provided estimates of climatic data for 1039 sites where species of Ramalina were found and 34 sites where no species of Ramalina were detected. Climatic variables were estimated from geographic location and elevation of each site using thin-plate splines fitted to average monthly climate data from meteorological stations (Leathwick & Stephens 1998; Leathwick 2001). Means and standard deviations for climatic variables were calculated for the sites in which a particular species of Ramalina was present, and species means provided the data for a principal components analysis using a centred correlation matrix (Kovach 1993). Ranges of climatic variables were subdivided into groups that were scaled either logarithmically (altitude) or linearly (temperature, precipitation, precipitation/evaporation ratios, solar radiation). Groups at the extremes of each range, where there were few records, were amalgamated. Percentage species frequency within each group was calculated from the ratio of the number of sites containing the species in question to the total number of sites within a particular group. Histograms showing species frequency with respect to annual means of minimum temperature and precipitation/evaporation (p/e) ratios, and altitude are included with graphs showing the geographic distribution of species. Species richness was determined by allocating records to 10-km squares on the New Zealand Map Grid (NZMG) and recording the number of species per square. Overall frequencies for a particular species were determined as the percentage of the recorded sites containing that species. Spread of species was measured as the distance (in km) across the longest axis of each species distribution, calculated

Bannister et al.—Ramalina in NZ


Fig. 1 Sites visited by the authors and associated collectors (see Acknowledgments) (O), and sites of herbarium specimens (Ø) with verified identifications of species of Ramalina (Blanchon et al. 1996, and subsequently).

123 Sites visited

Herbarium records

from the grid co-ordinates of the two most distant sites. The linear relationship between the logarithm of frequency and spread was used to compare the responses of sexually and asexually reproducing species. North and South Island records of R. canariensis are widely separated and are treated as two distinct groups for the purposes of analysis.

RESULTS AND DISCUSSION Recording and mapping Maps of species distribution must combine accurate species identification and location with adequate geographic coverage. The identity of herbarium specimens must be confirmed by inspection as not all herbarium specimens are correctly named. The revision of New Zealand species of Ramalina by Blanchon et al. (1996) enabled accurate identification of all the species of Ramalina found in New Zealand: previous herbarium records were checked and renamed where necessary (Blanchon 1994), as were herbarium specimens that came to light subsequently. There were obvious gaps in the maps of Ramalina distribution produced from herbarium and

other records collated by Blanchon et al. (1996), and there are still evident gaps in our more recent maps (Fig. 1). Gaps may be due to absence of a species or lack of collection. Species may be overlooked or not collected because they are too common, rare, or small, or because they grow in rarely visited or inaccessible areas. By targeting these gaps, the range of many species has been extended. Similar gaps in the distribution of vascular plants (e.g., western Taranaki, East Cape, inland Canterbury) were noted by Raven & Engelhorn (1971). In order to cover New Zealand, our recording was necessarily restricted to areas that could be readily accessed by roads (or, in western Fiordland and southern Stewart Island, by boat). Consequently, some mountainous or remote areas (e.g., the Kaikoura Ranges, regions to the west of the Southern Alps) remain under-recorded, although areas at higher altitudes (>1000 m) or with high precipitation/evaporation ratios rarely support species of Ramalina. The mode of recording also affects distribution maps. Individual herbarium records usually consist of single species collected from a particular site, whereas we searched for all species of Ramalina at any site that we visited. When records were grouped by site, only 27% of herbarium records recorded more than one spe-

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124

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to the Gisborne region (where it overlaps with the range ofR. glaucescens). Southern species There are four species that occur only in South Island: Ramalina inflexa, R.fimbriata, R. erumpens, andR. riparia (Fig. 8, 9). Herbarium records of R. inflexa (Blanchon et al. 1996) indicated a discontinuous distribution along coastal regions on the east coast of South Island from Cheviot to Invercargill. Our records (Fig. 8) extend its northern limit beyond Cheviot (Hundalee) and show that its distribution is continuous with extension further inland in the southern part of its range (sea level to 560 m). It is common at moderate altitudes, and absent from the driest and warmest areas, but is found over a greater range of temperatures and p/e ratios than the more restricted R. erumpens and R. riparia. Ramalina fimbriata (Fig. 8) is a rare species confined to Otago. It occurs under overhangs on schist tors at the highest altitudes (Galloway 2002), and on stream-side cliffs at moderately high altitudes, but also on coastal rocks and cliffs (Bannister 1998). It was previously considered to be confined to high altitudes (Blanchon et al. 1996), and is most frequent at high altitudes (Fig. 8), although most specimens were collected from sites below 1000 m. R.fimbriata is absent from the driest and wettest sites and tends to be most frequent in cooler locations (Fig. 8). Ramalina erumpens (Fig. 9) is an uncommon species, confined to coastal regions of South Island from Banks Peninsula to western Southland. We have extended its distribution within this range and identified herbarium specimens of R. banzarensis (CANU) from Bird Island, Foveaux Strait (Bannister & Blanchon 2002), and a previously unidentified Ramalina specimen from Banks Peninsula (CHR) as R. erumpens, and added them to our maps. Ramalina riparia (Fig. 9) shares a coastal distribution with R. erumpens from just north of Dunedin southwards. Within this range, R. riparia is more frequently encountered than R. erumpens. Like R. erumpens, R. riparia is most frequent at lower altitudes with moderate temperatures and rainfall. Habitat and substratum New Zealand species of Ramalina are mostly corticolous, and only R. fimbriata and R. meridionalis are predominantly saxicolous (Table 2). There are differences between our records of substrata and those from herbarium records. Herbarium records show proportionately greater occurrence of

133 some species on rock (R. canariensis, R. celastri, R. peruviana, R. unilateralis) and proportionately greater (R. peruviana) or lesser (R. celastri, R. glaucescens, R. inflexa) occurrence of some species on native trees and shrubs than in our records. This almost certainly relates to collection and recording. Our records were largely from roadside and inland sites and targeted areas where few collections had been made. These were biased towards native shrubs as these often proved to support species of Ramalina, whereas herbarium records were biased by the substantive collections that had been made from Auckland northwards in coastal areas and on islands which presented an extensive rocky shoreline. R. inflexa was most frequently recorded on exotic trees and this reflects its distribution in agricultural areas along the eastern coast of South Island in which there are very few native trees or shrubs. Likewise, records from wood (usually gates and fence posts) relate to agricultural areas with few native trees and shrubs, particularly in the south of South Island. Spread, range, and frequency The species of Ramalina that are established in New Zealand have three possible methods of propagation: vegetative spread by thallus fragmentation, dispersal of soredia, and sexual spread by ascospores (Bannister & Blanchon 2003). The establishment of a lichen from an ascospore requires that a

Table 2 Occurrence (% of records) of species of Ramalina on various substrata. Figures in bold indicate the type of substratum on which each species has been most frequently recorded. Woody plants Species

Rock

R.fimbriata R. meridionalis R. australiensis R. canariensis R. peruviana R. geniculata R. erumpens R. pacifica R. exigue lla R. unilateralis R. glaucescens R. riparia R. celastri R. inflexa

100 98

33 35 17 12 10 6 0 3 1 0 18 1

Native

Exotic

Wood

0 2 67 61 67 80 85 88 89 82 57 65 41 40

0 0 0 4 17 8 5 6 11 11 35 30 36

0 0 0 0 0 0 0 0 0 4 7 5 5 4

55

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135

Fig. 10 Frequency (%) of fertile (open circles) and vegetative (solid circles) species of Ramalina in New Zealand in relation to their spread (km).

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Spread (km) germinating spore forms a symbiotic relationship with a compatible alga (a species of Trebouxia in the case of Ramalina) whereas soredia are asexual reproductive structures consisting of algae surrounded by fungal hyphae. Consequently it is often considered that lichens producing soredia will be more successful in colonising new sites than those spread by ascospores alone. For example, Bowler & Rundel (1975) stated that "Propagation through vegetative diaspores appears to allow a higher survival rate, so that the frequency and abundance of plants using the asexual reproductive pathway is greater than that of species depending upon the germination of ascospores and a resynthesis of the symbiosis". Similar statements are often repeated in the literature, but seldom, if at all, tested quantitatively. Our distributional data gave us the opportunity to make a simple test. When log frequency is plotted against spread (Fig. 10), there is an overall linear relationship which is highly significant (F1,13 = 87.5, P < 0.001). The individual regressions for sexual and asexual species are also significant (P < 0.01). Analysis of covariance showed that the two regressions had significantly different elevations (P = 0.014) with the sexual species showing higher frequencies than asexual species at any particular level of spread. The two species with the highest frequencies and spread are R. celastri and R. glaucescens, both sexual species. R. unilateralis, a sorediate species, is the third most frequent and widespread

species. This is counter to the generally held view that sorediate species are more common and widespread than apothecial species. The British Isles have the same number ( 14) of species of Ramalina as New Zealand and distribution maps of British Lichen Society's database at Bradford University (M. R. D. Seaward pers. comm. 2002) show that, in contrast to New Zealand where many species have a discrete geographic range, the more common British species show wide latitudinal spread (often from Shetland to the Channel Islands). The most frequent species is Ramalina farinacea (sorediate), but the three next most frequent species (R. fastigiata, R. siliquosa, R.fraxinea) are apothecial, while the fifth and sixth most frequent species (R. subfarinacea, R. canariensis) are sorediate. As in the New Zealand examples, there is little evidence to suggest that British and Irish species of Ramalina with soredia are any more widespread or frequent than those producing ascospores. Species diversity and frequency High numbers of species of Ramalina are found in 10-km squares in the north-east and south-east of New Zealand, but R. celastri is the only frequent species in common. There are, however, areas where only R. celastri is found. Squares lacking any species of Ramalina tend to be associated with areas of high rainfall and/or high altitude or extreme temperatures (Fig. 2, 11).

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Fig. 11 Numbers of species ofRamalina for each 10-km square investigated in this study.


Bannister et al.—Ramalina in NZ Diversity of species of Ramalina is high in Northland and coastal South Otago. These areas are in parts of the country where there is a high degree of endemism in the vascular flora (Wardle 1991; McGlone et al. 2001), but the high degree of endemism found in north-west Nelson is not associated with high species diversity of species of Ramalina. Eight of the 14 species of Ramalina found in New Zealand are found in the coastal strip from Otago Peninsula southwards, and the Otago Peninsula has also been observed to have a high diversity of vascular plant species (Rogers & O verton 2000). Species diversity is inevitably related to habitat. Species of Ramalina are relatively light-demanding and are often found in relatively open habitats (e.g., in scrub, forest edges, and upper canopy). The most common species (e.g., R. celastri and R. glaucescens) are found in a wide range of habitats and on a variety of substrata and are often considered to be "weedy" species associated with anthropogenic disturbance. Other species are particularly associated with native forest remnants (R. erumpens, R. riparia, R. pacifica, R. geniculata), mangrove and coastal forest (R. exiguella), or native shrubs (R. unilateralis). Conservation of species diversity within Ramalina will depend on conservation of both native and exotic habitats (e.g., R. inflexa is most frequently found on exotic trees in an otherwise agricultural landscape on the east coast of South Island).

CONCLUSION Previous maps showing the distribution of species of New Zealand lichen genera include those for Pseudocyphellaria (Galloway 1988), Ramalina (Blanchon etal. 1996), and Sticta (Galloway 1997). The maps presented in this paper differ principally in their breadth and intensity of coverage. Accurate maps and locations have enabled us to make generalisations about the biodiversity, conservation status, and distribution of species of Ramalina in relation to geographic, habitat, and climatic factors. We hope that the results of this study will encourage others to follow suit, as accurate inventory not only tells us what is currently here but also provides a baseline for assessing future changes of species distribution.

137 ACKNOWLEDGMENTS We thank Barbara Anderson, Catherine Beard, Robyn Bridges, David Galloway, Jennifer Ide, Allison Knight, Kelvin Lloyd, Janice Lord, Keith Thompson, Isolde Ullmann, and Hugh Wilson for providing specimens from places that we had not visited, and M. R. D. Seaward for allowing us to use his unpublished distribution maps of British species of Ramalina.

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