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(Proteaceae) with Emphasis on Banksia Cuneata, a Rare and Endangered Species

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(Proteaceae) with Emphasis on Banksia Cuneata, a Rare and Endangered Species Heredity 79 (1997)394—40 1 Received 24 September 1996 Genetic diversity in Banksia and Dryandra (Proteaceae) with emphasis on Banksia cuneata, a rare and endangered species TINA L. MAGUIRE & MARGARET SEDGLEY* Department of Horticulture, Viticulture and Qenology, The University of Adelaide, Waite Campus, P/ant Research Centre, PMB 1, Glen Osmond, SA 5064, Australia Randomamplified polymorphic DNA (RAPD) markers were investigated as a tool for estimat- ing genetic diversity within 33 species of Banksia and three species of Diyandra. Three primers were used on DNA from 10 seeds per species, and band data were pooled to give between 52 and 89 bands per species, most of which were polymorphic. Genetic diversity was calculated using six published metrics on three species, for which allozyme data were also available. Based on between-method consistency, three metrics were chosen for analysis of the full data set. Levels of genetic diversity in Banksia and Diyandra ranged from 0.59 to 0.90. Based on this information, a detailed study was conducted on all 10 known populations of B. cuneata, a rare and endangered species, with a restricted geographical distribution in south-western Australia. Estimates of genetic diversity ranged from 0.65 to 0.74, which is high for a rare and endan- gered species. Analysis of molecular variance (AM0vA)wasused to partition RAPD variation within and between populations. Nearly all of the variation was attributable to individuals within populations, indicating a lack of population divergence. It is suggested that the combination of bird pollination and high outcrossing rates in B. cuneata maintain genetic diversity and cohesion between the populations. Keywords:AMOVA, Banksia,Diyandra, population genetics, Proteaceae, RAPDs. Introduction Banksia cuneata A. S. George is a rare and Scientificapproaches to the conservation and exploi- endangered species known only from 10 populations tation of plant genetic resources require a detailed (550 plants) in the central area of south-western knowledge of the amount and distribution of genetic Australia in an area of 90 km2. It is found in deep diversity within a species. Allozymes have been used yellow sands which occupy 10—15 per cent of the in Banksia to estimate genetic diversity and high area, giving it a fragmented distribution. Associated levels have been reported, amongst the highest with these soils is a rich and diverse flora dominated recorded for plants (Scott, 1980; Schemske & by species of the families Proteaceae, Myrtaceae and Lande, 1985; Carthew et al., 1988; Coates & Soko- Leguminosae. In the last 50—60 years land clearing for agriculture and other disturbances have reduced Iowski, 1992). More recently many workers have moved to DNA markers such as RAPDs, which have the B. cuneata population size to about 7 per cent of been useful for population genetic studies in a its original distribution, and it now occurs in number of genera (Chalmers et a!., 1992; Huff et a!., remnants of native vegetation. The mating system 1993). It has also been suggested that RAPDs may and patterns of genetic variation for six populations be an appropriate technique to monitor diversity in of B. cuneata have been determined using allozyme plant populations (Anderson & Fairbanks, 1990; electrophoresis (Coates & Sokolowski, 1992), and Virk el a!., 1995). estimates of outcrossing based on six loci ranged from 0.67—0.95, with low levels of selfing. The popu- lations were divided into two groups, with gene flow Correspondence. E-mail: [email protected] within groups but not between them. It was 394 1997The Genetical Society of Great Britain. GENETIC DIVERSITY IN BANKS/A 395 suggested that an ecological barrier to pollinator concentration was estimated by visual assessment of movement may be responsible for the restriction of band intensities, compared to salmon sperm between-group gene flow. genomic DNA standards. This study investigated levels of genetic diversity within species of Banksia and Diyandra using DNAamplification and documentation RAPDs, and evaluated six methods of data analysis. Based on this information, a detailed study deter- DNAamplification was performed in a MJ Research mining patterns of genetic variation within and Thermal Cycler. The programme commenced with a between all known populations of B. cuneata was denaturation step at 94°C for 5 mm, followed by 40 conducted. Ten populations were included in this cycles of 94°C for 1 mm, 36°C for 1 mm, 72°C for study, four of which had not been previously 2 mm, and terminated with an extension step at 72°C investigated. for 5 mm. Optimized reaction conditions were carried out in a 25 uL total volume containing Materials and methods 1 x Taq buffer (Gibco-BRL), 3 mrvi MgCl2, 200 M each of dNTP (dGTP, dATP, dCTP, dTAP), 1 unit Plant material of Taq polymerase (Gibco-BRL), 0.5 pL T4 gene 32 Seeds of 33 species of Banksia and three species of protein (Boehringher Mannheim), 1 M 10 mer Diyandra collected from natural populations were primer (Operon Technologies) and 10 ng template obtained from a commercial seed source (Ninde- DNA. Each reaction mix was overlaid with PCR thana Seed Service, WA). Seeds represent bulk grade paraffin oil. DNA amplification fragments collections from single or multiple sites of wild were separated by 2 per cent agarose gel (Seakem, populations. The species chosen represented the two Promega) electrophoresisusing TBE buffer sister genera Banksia and D,yandra, and two subge- (Sambrook et al., 1989). A negative control was nera, two sections and 13 series within the genus added in each run to test for contamination. In Banksia (George, 1981, 1988; Table 1). Ten seeds of order to test reproducibility, the selected primers each species were randomly selected for RAPD were tested three times on the same sample, for a random subset of three DNA samples. To aid inter- analysis. Banksia cuneata seed material was collected from pretation of band identity between gels, each all 10 known populations (Fig. 1). Twenty seeds contained a standard DNA sample and pGEM DNA were randomly selected, one per plant, from each of marker. Gels were stained with ethidium bromide four populations with in excess of 50 plants, and 10 and fragment patterns were photographed under seeds were collected, one per plant, from each of UV light with Polaroid 667 film. Polaroid photo- three populations with fewer than 50 plants. In three graphs were scanned using a transmission scanner cases, the populations comprised fewer than 10 (Hewlett Packard Scanjet IIcxJT). The intensity and plants, so a one-seed sample was taken from each. molecular weight of each visible band was deter- In total 125 plants, 25 per cent, of the remaining mined using the software CREAM (Kem-En-Tec Soft- 550 plants were sampled. ware Systems, Blue Sky Scientific). Based on the faintest visible band of the molecular weight marker (pGEM) a minimum intensity threshold was estab- DNAisolation lished. Bands with an intensity greater than the DNAwas extracted from each seed using a modifi- minimum threshold were scored as present. cation of the method of Weining & Langridge Sixty primers were evaluated for their suitability in (1991) comprising: phenol/chloroform/isopropanol a pilot survey (series OPA, OPB and OPC, Operon extraction for 5 mm on ice, DNA precipitation for Technologies). Three primers were selected for the 1 mm with ice-cold isopropanol and sodium acetate, initial study of genetic diversity in 33 species of with DNA recovered by centrifugation at 11 600g Banksia and three species of Diyandra. The primers for 10 mm. The pellet was washed twice with 70 per OPA-20 (GTTGCGATCC), OPB-03 (CATCC- cent ethanol, dried and dissolved in 50 tL of TE CCTG) and OPB-04 (GGACTGGAGT) gave repro- buffer, with 1.0 tL of RNAase (R40: 40 mg/mL ducible and informative markers. Five primers were RNAase A in TE), and stored at 4°C for up to selected for the detailed study on B. cuneata, 1 month. OPA-1, OPA-4, OPA-9, OPA-li and OPA-16 DNA was subjected to gel electrophoresis on 1.6 (Table 3). Band fragments included in the final per cent agarose gels in TBE buffer (Sambrook et analysis ranged between 2.5 kb and 100 bp in length a!., 1989), and stained with ethidium bromide. DNA (Fig. 2), were scored as present (1) or absent (0) for The Genetical Society of Great Britain, Heredity, 79, 394—401. 396 T. L. MAGU IRE & M. SEDGLEY Table1Banddata and estimates of diversity for 33 species of Banksia and three of Diyandra using RAPDs. Taxonomy follows George (1981, 1988) Number of bands Diversity (Standard deviation) Species PolymorphicMonomorphic TotalNei & Li (1979)Jaccard(1901)Russell & Rao (1940) Genus Banksia Subgenus Banksia Section Banksia Series Salicinae B. integnfolia * 82 0 82 0.77 (0.07) 0.74 (0.09) 0.86 (0.04) B. robur 61 0 61 0.79 (0.06) 0.77 (0.07) 0.88 (0.04) Series Grandes B. grandis 63 0 63 0.72 (0.06) 0.68 (0.07) 0.82 (0.03) B. solandri** 52 2 54 0.67 (0.07) 0.62 (0.09) 0.75 (0.05) Series Quercinae B. quercifolia** 51 1 52 0.73 (0.06) 0.69 (0.07) 0.83 (0.04) Series Bauerinae B. baueri 86 2 88 0.72 (0.06) 0.68 (0.08) 0.81 (0.05) Series Banksia B. baxteri 84 0 84 0.71 (0.06) 0.67 (0.07) 0.82 (0.03) B. candolleana 60 0 60 0.74 (0.06) 0.71 (0.08) 0.83 (0.04) B. menziesü 79 5 84 0.65 (0.06) 0.59 (0.07) 0.77 (0.04) B, serrata 53 1 54 0.73 (0.07) 0.70 (0.08) 0.84 (0,04) Series Crocinae B.
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