Available online at www.sciencedirect.com R Review of Palaeobotany and Palynology 124 (2003) 113^129 www.elsevier.com/locate/revpalbo Pollen distribution in marine surface sediments o¡shore Western Australia Sander van der Kaars a; , Patrick De Deckker b a Centre for Palynology and Palaeoecology, School of Geography and Environmental Science, Monash University, Clayton, VIC 3800, Australia b Department of Geology, The Australian National University, Canberra, ACT 0200, Australia Received 1 June 2001; accepted 25 November 2002 Abstract We have examined the pollen content in sediments from the top of 38 cores taken offshore Western Australia (WA). Water depth for these cores ranges between 81 and 4090 m. All samples are used to plot maps of total pollen and total Pteridophyta spore concentration. Only the 26 core tops that yielded sufficient pollen grains ( s 35) are used in the present study to plot percentage maps for individual pollen taxa, including that of the genus Pinus (recently introduced to Australia). Five major bioclimatic zones are recognised by the distribution of pollen assemblages offshore WA. These are related to the amount and seasonality of rainfall and vegetation distribution on land. They reflect: (1) the open Eucalyptus forests and woodlands of northern WA with high and summer rainfall; (2) the open Eucalyptus woodlands with Acacia and grasslands of northwestern WA with lower (400^600 mm) and summer rainfall; (3) the open Acacia shrublands with grasses and grasslands of northwestern WA with low (300^400 mm) and summer rainfall; (4) the open Acacia shrublands of central WA with low ( 6 300 mm) and summer as well as winter rainfall; and (5) the open Eucalyptus forests and mixed shrublands of southwestern WA with lower (300^600 mm) and winter rainfall. The influence of Indonesian-derived waters via surficial currents such as the Leeuwin Current is recognised by the presence of Pteridophyta spores in the sediments at northern sites. ; 2003 Elsevier Science B.V. All rights reserved. Keywords: pollen; marine palynology; Pteridophyta spores; Western Australia; core tops 1. Introduction Palynological records from relatively dry areas are di⁄cult to obtain by traditional terrestrial studies due to the paucity of lakes and/or bogs in these areas. However, marine palynological re- * Corresponding author. E-mail address: [email protected] (S. van der Kaars). search can provide important information on continental vegetation and climatic developments, as well as the direction of predominant wind and ocean currents and represent an alternative source of palaeoecological information for some of these relatively dry areas (for instance, West Africa, northwestern Australia and the Arabian Sea area). It has been shown that detailed palaeoecological interpretations are possible for these regions by comparison of core top samples and marine core spectra (see Hooghiemstra, 1988; 0034-6667 / 03 / $ ^ see front matter ; 2003 Elsevier Science B.V. All rights reserved. doi:10.1016/S0034-6667(02)00250-6 PALBO 2515 17-3-03 114 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 Hooghiemstra and Agwu (1988)). This approach is particularly appropriate for o¡shore Western Australia (WA) as there are very few records on land (see Wallis, 2001 and references therein), especially in the northern regions of this very large arid Australian state. Despite the importance of marine palynological records in the palaeoecological study of WA (van der Kaars, 1991; Wang et al., 1999; van der Kaars and De Deckker, 2002), very little is known about the recent pollen distribution patterns in marine sediments from the waters of WA. This study aims to provide a basis for re¢ned interpretation of the marine palynological records of WA through analysis of pollen assemblages from marine core top samples along its coast. The pollen distribution in marine surface sediments from the waters of WA is analysed in relation to the onshore vegetation distribution, climate and pollen transport by water and/or wind. The results will be of use for the bioclimatic interpretation of palynological records and as a guide in the selection of cores and future coring sites for marine palynological research in the region. Marine palynological studies are proving very important nowadays when attempting to reconstruct palaeoclimates, and even more so when aiming at identifying links between signals at sea and on land. Such studies can only be achieved through better knowledge of the distribution of pollen at sea before interpreting the fossil record. This is exactly what our present work aims at: providing a sound base for palaeoecological studies of an arid region such as a large part of WA. 2. Environmental setting WA encompasses approximately 20 degrees of latitude and therefore covers a wide variety of climatic conditions. The northern part of WA comprises the humido^arid tropical summer rain region, the centre the subtropical arid region and the southern part the arido^humid winter rain region (Sturman and Tapper, 1996). Annual precipitation along the transect varies from between 400 and 1600 mm in the north and south to less than 300 mm in the centre (see Fig. 1). In the north, rainfall is predominantly produced in summer and in the south rainfall is predominantly a winter feature, while the area in between receives both summer and winter rain (see Fig. 1). The vegetation along a north^south transect varies from open Eucalyptus forests with a grassy understorey and grass-rich open Eucalyptus woodlands in the north, to grass-rich open Acacia shrublands and (coastal) grasslands, grading to open Acacia shrublands in the centre and to mixed shrublands, grading to open Eucalyptus woodlands and Eucalyptus forests in the south (see Fig. 2). In general, the vegetation north of latitude 23‡S is much richer in grasses than the area to the south. The area of interest here, i.e. the eastern Indian Ocean and WA, is characterised by contrasting seasons that are re£ected by changes in ocean surface current directions. The strengths and directions of these surface currents are strongly controlled by winds. The generalised surface circulation in summer is characterised by predominantly northwesterly winds in northern Australia that are accompanied by much rainfall, as part of cyclonic depressions. The region registers a complete change of wind direction to southeasterly during the austral winter. At that time, the northern part of WA is dry and the southern portion of the state is wet. Up to 50% of the rainfall in northern WA can be delivered by cyclones in summer (Wyrwoll, 1993). These cyclonic depressions cause strong river discharge, and it is likely that much pollen is transported o¡shore during that time. Ocean currents are also likely to in£uence the distribution of pollen. The major current that follows the periphery of WA is the Leeuwin Current (LC), which is an o¡shoot of the Indonesian Through£ow (Cresswell, 1991; Wij¡els et al., 1996). The LC is characteristically a low salinity, warm water current that £ows poleward as far as Cape Leeuwin; it eventually engenders large eddies of low density water at higher latitudes in the vicinity of Geraldton down to Cape Leeuwin (see Cresswell, 1991 for more details). The strength of the LC is strongest during the winter season. The cold, equatorward-£owing West Australian Current travels below the LC. It is found along the west coast of WA north- PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 115 Fig. 1. Map showing the core top locations, topographic names mentioned in the text and rainfall data (source: Commonwealth of Australia, Bureau of Meteorology, 2000, http://www.bom.gov.au). wards to North West Cape from where it migrates west to form part of the South Equatorial Current. This current has the potential to transport pollen from the southern portion of WA northward. 3. Materials and methods Selected core tops obtained during two RV Franklin cruises, Fr10/95 (December 1995) and Fr2/96 (February^March 1996), in the waters of PALBO 2515 17-3-03 116 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 Fig. 2. Map showing core top location and relevant vegetation of WA (simpli¢ed after AUSLIG, 1990). Heavy solid line indicates extent of sand dune country. WA were sampled for this study. The two cruises aimed at gathering oceanographic as well as palaeoceanographic data, which included the collection of some 52 deep-sea sediment cores along a depth transect from 81 to 4090 m. The same core tops have been used to study the distribution of planktonic foraminifera (Mart|¤nez et al., 1998), terrigenous clays (Gingele et al., 2001) and benthic foraminifera (Murgese, submitted). Samples for pollen analysis were prepared using PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 the following method. The sediment was suspended in about 40 ml of tetra^sodium^pyrophosphate ( P 10%), sieved over 210- and 7-Wm meshes, followed by hydrochloric acid (10%) treatment and heavy liquid separation (sodium^polytungstate, SG 2.0, 20 min at 2000 rpm, twice) and hydro£uoric acid (50%) treatment. The organic residue was stained with safranin. Slides were mounted in glycerol and sealed with para⁄n 117 wax. A known amount of Lycopodium marker spores was added to each sample prior to chemical treatment in order to establish palynomorph concentrations. All slides were counted along evenly spaced transects using a Zeiss Axioskop microscope at U630 magni¢cation. All percentage values were calculated on the total pollen sum (all pollen counted, excluding unknown pollen grains and Pteridophyta spores). Table 1 Core top samples Fr10/95 and Fr2/96 cruises Core Location Fr95-2 Fr95-3 Fr95-4 Fr95-5 Fr95-6 Fr95-7 Fr95-8 Fr95-9 Fr95-10 Fr95-11 Fr95-12 Fr95-13 Fr95-14 Fr95-15 Fr95-16 Fr95-17 Fr95-18 Fr95-19 Fr95-20 Fr95-25 Fr95-26 Fr95-27 Fr96-4 Fr96-6 Fr96-7 Fr96-11 Fr96-12 Fr96-14 Fr96-17 Fr96-19 Fr96-21 Fr96-22 Fr96-23 Fr96-24 Fr96-25 Fr96-26 Fr96-28 Fr96-29 12‡32.86PS, 126‡14.84PE 13‡14.53PS, 124‡00.2PE 13‡55.18PS, 122‡01.51PE 14‡00.55PS, 121‡01.58PE 14‡19.67PS, 121‡09.81PE 14‡42.58PS, 120‡32.74PE 14‡54.97PS, 120‡57.49PE 18‡07.63PS, 118‡00.92PE 18‡08.93PS, 116‡01.29PE 17‡38.57PS, 114‡59.93PE 18‡14.7PS, 114‡59.63PE 18‡49.26PS, 113‡58.26PE 20‡02.71PS, 112‡39.73PE 19‡53.75PS, 112‡13.37PE 20‡59.83PS, 112‡59.35PE 22‡07.74PS, 113‡30.11PE 22‡59.64PS, 112‡49.86PE 24‡14.11PS, 110‡00.18PE 24‡44.67PS, 111‡49.75PE 28‡43.93PS, 113‡22.08PE 29‡14.42PS, 113‡33.48PE 30‡30.14PS, 114‡16.64PE 28‡43.02PS, 113‡23.32PE 28‡25.21PS, 112‡17.37PE 26‡58.76PS, 111‡20.13PE 23‡57.16PS, 108‡22.14PE 23‡44.23PS, 108‡31.91PE 19‡24.64PS, 110‡30.40PE 12‡14.80PS, 112‡44.27PE 12‡22.76PS, 114‡16.96PE 14‡48.68PS, 114‡16.37PE 16‡34.71PS, 113‡11.98PE 16‡54.81PS, 113‡20.14PE 16‡55.61PS, 114‡15.46PE 16‡54.65PS, 115‡15.90PE 16‡54PS, 115‡31PE 18‡47.93PS, 116‡20.23PE 18‡57.81PS, 116‡23.52PE a Water depth (m) 81 182 470 2472 2177 1445 678 498 1462 2458 2034 1454 997 1393 1211 1093 1055 1974 841 1010 1738 843 936 3575 3090 2404 2100 4090 2571 3355 2919 2501 1967 1603 1666 1958 502 344 Sample dry weight (g) Pollen counta Spore count 3.44 3.50 1.76 4.00 3.70 2.16 4.25 3.78 3.78 3.55 2.53 3.54 3.88 4.88 3.33 4.36 3.04 3.82 2.00 3.92 4.75 5.53 3.15 2.94 2.27 2.17 3.77 3.15 3.56 2.36 6.09 4.19 3.80 5.73 3.97 4.08 3.92 4.78 109 45 115 114 118 147 156 173 122 43 44 100 153 151 122 196 126 9 105 117 134 108 94 2 37 5 1 3 23 5 2 4 2 4 28 39 244 295 ^ ^ 2 27 56 8 5 ^ 10 6 6 4 2 1 5 ^ 2 2 1 5 7 3 2 ^ 3 ^ ^ 1 30 12 1 ^ ^ ^ 3 1 1 1 Excluding unknown grains. PALBO 2515 17-3-03 Pollen concentration (pollen/g) 104 109 638 239 81 1195 1026 805 70 24 38 54 369 165 106 1188 309 15 112 147 101 99 170 3 35 11 1 4 15 10 3 5 5 7 19 23 719 1005 Spore concentration (spores/g) ^ ^ 11 57 39 65 33 ^ 6 3 5 2 5 1 4 ^ 5 3 1 6 5 3 4 ^ 3 ^ ^ 1 20 23 1 ^ ^ ^ 2 1 3 3 118 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 4. Results In all 38 core top samples were processed for this study (see Table 1). All samples were used to establish pollen and Pteridophyta spore concentrations. However, any sample with a pollen count of less than 35 pollen grains was omitted from further analysis (12 samples in total). The remaining 26 samples were used to establish the pollen distribution patterns. Samples Fr95-3, Fr95-11, Fr95-12, Fr96-7, and Fr96-26 have low pollen sums, between 37 and 45 pollen grains, and their percentage values can be considered as less reliable. Therefore, percentage values obtained from these ¢ve samples were only used in the description of the pollen assemblages when they agreed with the percentage values obtained from nearby samples with higher pollen sums. Selected results are presented as isopoll (i.e. equal pollen percentage value) maps (Figs. 3^6), while the complete results are presented as a pollen percentage diagram in Fig. 7. In the pollen diagram the core locations are arranged in north^south direction according to their latitude. 4.1. Pollen and Pteridophyta concentrations The pollen concentrations show a distinct pattern that roughly follows the outline of the continental shelf (see Fig. 3A). The pollen concentration values decrease rapidly with increasing distance from the shore. The northwestern part of the WA waters shows higher pollen concentrations than their southwestern portion, with a relatively broad band of values above 1000 pollen grains per gram being restricted to the northwest. Values between 100 and 1000 pollen grains per gram occur in a narrow area within the northwestern sector and the area is located further from shore than in the southwest. Pteridophyta spore concentrations are much lower, in general below ¢ve spores per gram at all locations except for the most northern ones (Fig. 3B). There, concentration values of more than ten spores per 119 gram occur, and it appears that Pteridophyta spore concentration values increase with increasing distance from shore. 4.2. Rhizophoraceae Rhizophoraceae (Fig. 3C) occur only with low percentages or are absent altogether at most sites, except for the northern part of the waters of WA where higher values (up to 3.5%) are encountered. 4.3. Herbs Asteraceae Tubuli£orae (Fig. 3D) show a gradual increase from the northern waters to approximately 25‡S, from less than 5% to more than 20%. Further south values vary, but are generally higher than 20%. The Chenopodiaceae/Amaranthaceae percentages (Fig. 4A) also show a gradual increase. From the northern waters to approximately 23‡S, values increase from less than 5% to more than 20%. Further south, percentages gradually decrease to less than 10%. The percentages for Poaceae (Fig. 4B), on the other hand, decrease southward. Values greater than 40% occur in the northern part of the waters of WA, decreasing to less 10% in the southern part. The Cyperaceae percentages (Fig. 4C) vary more but are, in general, higher o¡shore the northern part of WA, between 5 and 10%, and lower southward, mostly below 5%. Restionaceae (Fig. 4D) occur in low numbers north of latitude 16‡S, are absent between approximately latitudes 16 and 20‡S, and reach somewhat higher values southward, with the highest ^ more than 10% ^ at the southernmost site. 4.4. Trees and shrubs The distribution pattern of Callitris (Fig. 5A) is patchy and pollen representation is poor. Only north of latitude 16‡S, around latitude 20‡S as Fig. 3. Isopoll maps for pollen (A) and Pteridophyta spore (B) concentrations, as well as Rhizophoraceae (C) and Asteraceae Tubuli£orae (D) percentages, in relation to the onshore vegetation. Open circles indicate absence. PALBO 2515 17-3-03 120 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 well as around latitude 30‡S Callitris pollen is recorded with low percentages. Gyrostemon pollen (Fig. 5B) is present with values of around 5% at most locations. However, around latitude 19‡S, values of more than 10% are found. Acacia pollen (Fig. 5C) is recorded with low percentages only, generally around 1%, north of 19‡S and between latitudes 20 and 29‡S. High values for Myrtaceae pollen (Fig. 5D), greater than 10%, occur north of latitude 16‡S and south of latitude 28‡S. Values in between the two areas are below 10%, with values of less than 5% recorded between latitudes 16 and 20‡S. Casuarina (Fig. 6A) is absent at latitude 18‡S; it is present in low abundance, around 1%, at most other locations. Only between latitudes 26 and 28‡S values greater than 5% are found. The introduced genus Pinus (Fig. 6B) is present, but with a patchy distribution north of 25‡S. South of that latitude Pinus pollen is well represented, and values greater than 10% are recorded at the two southernmost sites. 121 cladus, Podocarpus) and the exotic taxa (Alnus and Pinus). The other zones have 29 taxa or less. 4.5.2. Zone B Between 16 and 18‡30PS, the pollen assemblages are characterised by the absence of Rhizophoraceae, low Asteraceae Tubuli£orae ( P 6%), high Poaceae ( P 50%), high Chenopodiaceae/Amaranthaceae (10^17%), and varying Cyperaceae ( P 7%) values, the absence of Restionaceae, a virtual absence of Callitris, varying Gyrostemon (6^14%), low Acacia ( P 2%), varying Myrtaceae (1^8%), and low Casuarina values (0^3%). 4.5.3. Zone C Characteristic of the pollen assemblages between 18‡30PS and 21‡S are the virtual absence of Rhizophoraceae, high Asteraceae Tubuli£orae (10^20%), high Poaceae (30^50%), high Chenopodiaceae/Amaranthaceae (6^30%), varying Cyperaceae (1^4%), low Restionaceae (0^2%), low Callitris (0^1%), varying Gyrostemon (1^14%), varying Acacia (0^3%), varying Myrtaceae (3^ 9%) and low Casuarina values (0^2%). 4.5. Pollen diagram The pollen diagram (Fig. 7) has been divided into ¢ve zones. Each zone is brie£y described using only the 11 most common taxa. 4.5.1. Zone A The pollen assemblages between 12 and 16‡S are characterised by a low presence of Rhizophoraceae (0^4%), low Asteraceae Tubuli£orae (1^ 5%), high Poaceae ( P 60%), low Chenopodiaceae/Amaranthaceae ( 6 5%), and high Cyperaceae values (5^14%), low presence of Restionaceae, the presence of Callitris ( P 2%), varying presence of Gyrostemon ( P 5%), varying presence of Acacia (1^5%), high abundance of Myrtaceae (10^20%), and low presence of Casuarina (0^4%). These assemblages also show the highest diversity, with more than 44 taxa not including the Indonesian taxa (Dipterocarpaceae, Dacrycarpus, Phyllo- 4.5.4. Zone D The pollen assemblages between latitudes 21 and 26‡S are characterised by a virtual absence of Rhizophoraceae, high Asteraceae Tubuli£orae (30^38%), lower Poaceae (11^25%), high Chenopodiaceae/Amaranthaceae (14^33%), varying Cyperaceae (1^4%), and low Restionaceae (0^2%), the absence of Callitris, low Gyrostemon (2^3%), varying Acacia (1^5%), low Myrtaceae (7^9%), and low Casuarina values (1^2%). 4.5.5. Zone E Between 26 and 31‡S, the pollen assemblages are characterised by the absence of Rhizophoraceae, high Asteraceae Tubuli£orae (18^32%), varying Poaceae (2^14%), lower Chenopodiaceae/Amaranthaceae (4^9%), low Cyperaceae (2^ 4%), higher Restionaceae (1^11%), low Callitris (0^1%), higher Gyrostemon (3^7%), low Acacia Fig. 4. Isopoll maps for Chenopodiaceae/Amaranthaceae (A), Poaceae (B), Cyperaceae (C), and Restionaceae (D) percentages, in relation to the onshore vegetation. Open circles indicate absence. PALBO 2515 17-3-03 122 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 123 Fig. 6. Isopoll maps for Casuarina (A) and Pinus (B) percentages, in relation to the onshore vegetation. Open circles indicate absence. (0^3%), higher Myrtaceae (8^33%), and high Casuarina values (2^5%). A simple correspondence analysis using all of these common taxa except Rhizophoraceae seems to support this zonation, as sites for each zone are plotted in distinct ¢elds that do not show any overlap (Fig. 8). The ¢rst four ordination axes in the correspondence analysis had the following eigenvalues: Axis 1, eigenvalue 0.218; Axis 2, eigenvalue 0.130; Axis 3, eigenvalue 0.055; Axis 4, eigenvalue 0.037. According to ter Braak (1987), only ordination axes with an eigenvalue greater than 0.3 are considered to explain major variations in the data. We therefore assume that for this data set the ¢rst four ordination axes do not explain major variations in the pollen assemblages. As a result, the following discussion makes largely use of information obtained from the isopoll maps and the pollen diagram. 5. Discussion Newsome (1999) found a good relationship between the patterns of vegetation and pollen distribution onshore in the semi-arid southwestern part of WA. Our results indicate that the onshore distribution of the present-day vegetation is re£ected also by the pollen distribution in the marine sediments o¡shore WA. Vegetation types such as the northern and southern Eucalyptus forests can easily be recognised by their pollen signal in the marine sediments, with Myrtaceae percentages of around 20% or higher. The latter can be Fig. 5. Isopoll maps for Callitris (A), Gyrostemon (B), Acacia (C), and Myrtaceae (D) percentages, in relation to the onshore vegetation. Open circles indicate absence. PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 Fig. 7. Pollen diagram showing percentage values for individual taxa from core tops o¡shore WA. 124 PALBO 2515 17-3-03 Fig. 7 (Continued). S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 PALBO 2515 17-3-03 125 126 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 0.8 Fr95-2 0.6 Fr95-27 ♦ >900 mm, summer rain Fr95-26 ♦ Fr95-4 ♦ 0.4 Fr95-3 ♦ Axis 2 0.2 ♦ ♦ Fr96-4 ♦ Fr96-7 ♦ Fr95-8 Fr95-6 ♦ 300-600 mm, winter rain Fr95-7 ♦ Fr95-5 0 ♦Fr95-25 ♦ Fr95-11 ♦ ♦ Fr95-13 400-600 mm, summer rain Fr95-10 -0.2 Fr95-12 -0.4 ♦ ♦ Fr95-16 ♦ Fr96-26 ♦ ♦ ♦ ♦ Fr95-14 Fr95-15 ♦ Fr95-20 ♦ Fr95-18 ♦ Fr96-28 Fr95-9 300-400 mm, summer rain <300 mm ♦ Fr96-29 -0.6 ♦ Fr95-17 -0.8 -1 -0.8 -0.6 -0.4 -0.2 0 0.2 0.4 0.6 0.8 1 1.2 Axis 1 Fig. 8. Correspondence plot for the core tops o¡shore WA, using ten of the most common taxa only. separated from each other by high Poaceae percentages in the marine pollen assemblages derived from the northern forests and by high Asteraceae Tubuli£orae percentages in the marine pollen assemblages derived from the southern forests. The grass-rich vegetation types north of latitude 23‡S, from the area of predominantly summer rain, are re£ected by high Poaceae percentages in the marine sediments north of that latitude. The poor representation of Acacia pollen in the marine sediments makes the recognition of the pollen signal derived from open Acacia shrublands di⁄cult. The Acacia shrublands, with grasses north of latitude 23‡S, may be recognised by high Asteraceae Tubuli£orae, Poaceae and Chenopodiaceae/ Amaranthaceae percentages, while those poor in grasses ^ south of latitude 23‡S ^ have higher percentages of Asteraceae Tubuli£orae and Chenopodiaceae/Amaranthaceae, and distinctly lower ones of Poaceae. For most of the taxa presented in the isopoll maps, the distribution in marine sediments seems to re£ect their onshore distribution. For instance, the disjunct distribution of Callitris in the marine sediments seems to re£ect the distribution of Callitris onshore, with Callitris intratropica being restricted to the northern part of the state and Callitris glaucophylla to the central arid area and (topographically high) ranges, and with C. verrucosa, C. preissii, C. roei and C. drummondii in the southwest of WA (see Fig. 9). Casuarina occurs in high percentages only in marine sediments adjacent to the Casuarina shrublands and Casuarina mixed shrublands in southwestern WA. Rhizophoraceae are best represented in the marine sediments of the northern part of the state, adjacent to where the most extensive mangrove vegetation in WA is found. Even the introduced Pinus shows its highest values (more than 10%) in the waters southwest of WA, close to the pine plantations around Perth (see AUSLIG, 1990). The distribution of Restionaceae is restricted to the southwest of WA (see Fig. 10) south of latitude 23‡S; this is well re£ected by the pollen distribution in the marine sediments, with the exception of the presence of Restionaceae pollen in the marine sediments PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 127 north of 16‡S. However, pollen from Centrolepidaceae and Restionaceae look very similar and Centrolepidaceae do occur in the northern part of the state (see Fig. 11). It is therefore likely that the ‘Restionaceae’ pollen grains north of 16‡S were actually derived from Centrolepidaceae. The increase in Pteridophyta spore concentration, with increasing distance from shore, especially in the northwest of the WA waters, probably re£ects an in£ux of spores transported from the Indonesian region into the waters of WA via the LC. Pteridophyta form an important element of the ever-wet rainforests of Indonesia and Pteridophyta spores are transported easily by water and ocean currents (van der Kaars, 2001). With increasing distance from shore, the composition of the marine pollen assemblages is clearly in£uenced by di¡erential pollen transport and longshore currents. For instance, relative Fig. 10. Distribution of Restionaceae in WA (after Hnatiuk, 1990). abundance of Chenopodiaceae/Amaranthaceae seems to decrease with increasing distance from the shoreline, while the Poaceae values do the opposite ; they increase with increasing distance from shore. The pollen distribution pattern seems to form a coherent pattern. Even with the limited number of available samples (38), it is possible to present a tentative correlation between the marine pollen distribution in the waters of WA and the onshore vegetation and climate. Table 2 presents the proposed relationship between the major bioclimatic zones in WA and the main components of the marine pollen assemblages (see also Fig. 8). 6. Conclusions Fig. 9. Distribution of Callitris intratropica and C. glaucophylla in WA (after Hnatiuk, 1990; Bowman and Harris, 1995). Our pollen concentration map indicates that suitable core sites for marine palynological re- PALBO 2515 17-3-03 128 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 Fig. 11. Distribution of Centrolepidaceae in WA (after Hnatiuk, 1990). search o¡shore WA could be found in a relatively narrow band parallel to the coastline up to 400 km o¡shore in the northwest and 150 km in the southwest of WA. Suitable sites are mostly located at water depths ranging between 400 and 2500 m. In general, the bioclimatic zones are well re£ected in the marine pollen assemblages o¡shore WA. This phenomenon may prove to be a useful tool for the interpretation of the Late Quaternary marine palynological records from the region. The area of WA, with predominantly summer rain, can be recognised in the marine pollen assemblages north of latitude 23‡S by Poaceae percentages s 30. The area of predominantly winter rain is re£ected in the marine pollen assemblages south of latitude 26‡S by Poaceae percentages between 10 and 20. The area with low summer and winter rainfall between 21 and 26‡S is re£ected by Asteraceae Tubuli£orae percentages s 30. The arid central area of WA is re£ected in the marine pollen assemblages between 18‡30PS and 26‡S by Chenopodiaceae/Amaranthaceae and Asteraceae Tubuli£orae percentages both v 10, with combined percentages v 30, and Myrtaceae percentages 6 10. At this stage, we are unable to decipher whether the pollen distribution in the marine sedi- Table 2 Proposed relationship between bioclimatic zones in WA and the composition of marine pollen assemblages Bioclimatic zones Myrtaceae Poaceae Cyperaceae Asteraceae Tubuli£orae (%) (%) (%) (%) Chenopodiaceae/ Amaranthaceae (%) Northern WA: grass-rich open Eucalyptus forests and open s 10 Eucalyptus woodlands; annual precipitation s 900 mm, predominantly summer rain 40^60 5^10 65 6 10 Northwestern WA: open Eucalyptus woodlands with Acacia, 6 10 and grasslands; annual precipitation 400^600 mm, predominantly summer rain P 50 P5 P5 10^20 Northwestern WA: open Acacia shrublands with grasses and grasslands; annual precipitation 300^400 mm, predominantly summer rain 30^50 65 10^20 10^30 Central WA: open Acacia shrublands; annual precipitation 5^10 6 300 mm 6 25 65 30^40 15^35 Southwestern WA: open Eucalyptus forests and mixed shrublands; annual precipitation 300^600 mm, predominantly winter rain 10^20 65 20^30 6 10 6 10 s 10 PALBO 2515 17-3-03 S. van der Kaars, P. De Deckker / Review of Palaeobotany and Palynology 124 (2003) 113^129 ments is principally caused by aeolian or by £uvial processes. We expect that close to shore in the northern regions of WA, where rivers contribute to a large in£ux of freshwater during the summer rainy season, pollen are principally transported by water as well as wind. The pollen distribution further o¡shore is likely to be predominantly of aeolian origin. The LC, which follows the outline of WA and partly originates from the Indonesian Through£ow, is more than likely the ‘carrier’ of Pteridophyta spores of Indonesian origin. Acknowledgements We would like to thank Gary Swinton for drafting the maps. This research was supported by a Monash Logan Research Fellowship to S.v.d.K. The cores were obtained by P.D.D. with help of several ARC grants and access to the Australian national facility RV Franklin. We thank the reviewers for their many useful comments. References AUSLIG, 1990. Vegetation. 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