de Kock.qxd Freshwater mollusc diversity in the Kruger National Park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall K.N. DE KOCK, C.T. WOLMARANS and L.H. DU PREEZ De Kock, K.N., C.T. Wolmarans and L.H. du Preez. 2002. Freshwater mollusc diversi- ty in the Kruger National Park: a comparison between a period of prolonged drought and a period of exceptionally high rainfall. Koedoe 45(2): 1-11. Pretoria. ISSN 0075-6458. Most of the previous records of the freshwater molluscs from the Kruger National Park date back to 1966 and earlier. On account of several droughts between 1966 and 1995, a survey was done in 1995 to evaluate the effect of these droughts on the mollusc pop- ulation. As a result of extensive rainfall between 1995 and 2000 another survey was con- ducted to establish the effect of a period of exceptionally high rainfall on the species’ diversity of the mollusc population. With the exception of three habitats, an increase in mollusc diversity was recorded for all the other habitats previously surveyed. One of the invader freshwater snail species, Aplexa marmorata, collected in only one habitat in 1995, was found in seven habitats located mainly in the south during the 2001 survey. Another interesting finding was that juvenile specimens of Lanistes ovum, of which large specimens were present prior to 1966 but none found in 1995, were present in the Sirheni Dam in 2001. From this study the positive effect of the high rainfall on the species’ diversity is highly evident. Key words: Mollusca, freshwater molluscs, invader species, rainfall, Kruger National Park, South Africa. K.N. de Kock, C.T. Wolmarans and L.H. du Preez, School of Environmental Sciences and Development (Zoology), Potchefstroom University for Christian Higher Education, Private Bag X6001, Potchefstroom 2520, Republic of South Africa. ISSN 0075-6458 1 Koedoe 45/2 (2002) Introduction The importance of regular, normal rainfall to maintain the bio-diversity in conservation areas in South Africa cannot be over-empha- sized, especially for those aquatic organisms which lack the ability to migrate to other habitats during periods of serious drought. This inability to migrate leads to the situation where re-population of isolated habitats, such as dams and pans, is largely dependent on time and passive dispersal of biota. Cer- tain freshwater invertebrates, however, have the ability to aestivate, a strategy enabling them to survive prolonged periods of drought. Apart from these freshwater snail species, among others, Bulinus reticulatus Mandahl-Barth, B. forskalii (Ehrenberg, 1831), B. tropicus (Krauss, 1848) and the B. africanus group of species (Brown 1994), it is mostly the eggs of some freshwater crus- taceans such as Triops, Eocyzicus and Branchipodopsis which can survive periods of drought (Davies & Day 1997). With regard to the freshwater molluscs, it is known that the largest bio-diversity is to be found in the tropical and sub-tropical areas of the world. In South Africa it is mainly the Limpopo, Mpumalanga and KwaZulu-Natal provinces that represent the tropical and sub- tropical regions with the Kruger National Park situated in both the first two mentioned provinces. A large variety of freshwater mol- luscs collected up to 1966 were reported by Oberholzer & Van Eeden (1967) for the Kruger National Park. de Kock.qxd 2005/12/09 10:01 Page 1 Koedoe 45/2 (2002) 2 ISSN 0075-6458 The distribution of freshwater molluscs is not a static process and is mainly influ- enced by the availability of suitable water habitats. The most recent account of the diversity of freshwater molluscs in the park, prior to the 2001 survey, dates back to 1995 and is a report by De Kock & Wol- marans (1998) on a survey done after the drought of 1991-1992. Between 1966 and 1995, the park was struck by several droughts, namely those in 1982-1983 and 1986-1987—the former also being the severest—prior to the third drought of 1991-1992. According to Zambatis & Biggs (1995) several of the rainfall moni- toring stations recorded the lowest rainfall in history. Between 1995 and 2001, how- ever, average to above normal high rainfall figures were recorded for most of the rain- fall monitoring stations. This situation prompted the idea to do a follow-up of the 1995 survey to evaluate the effect of the availability of more permanent water habi- tats on the freshwater mollusc diversity, and to compare results with the 1995 sur- vey. Methods and materials During the present investigation we concentrated mainly on water bodies that had been surveyed for snails during the 1995 collection. We also concen- trated on several other water bodies that were dry during the 1995 survey, or seemed suitable for supporting molluscs but of which there were no previous records in the National Freshwater Snail Collection (NFSC). A total of 43 habitats were surveyed during 2001. In each survey, all the habi- tat types were screened for molluscs by means of aluminium mollusc scoops. The collected mol- luscs were identified by the senior author who had mainly been responsible for the identification of the molluscs in the NFSC since 1973. The mol- luscs were subsequently incorporated into the National Freshwater Snail Collection. Habitats without surface water were screened for mollusc shells. Rainfall records, for the periods 1982-1983, 1985-1987, 1991-1994 and 1995- 2001, were obtained from the Kruger National Park for the monitoring stations nearest to the habitats surveyed. Fig. 1. Water bodies (•) sampled during the 2001 survey. Rainfall monitoring stations (♦). de Kock.qxd 2005/12/09 10:01 Page 2 Results Water bodies sampled during the 2001 sur- vey are depicted in Fig. 1. The results of this investigation are presented in Tables 1, 2 & 3 and Figs. 2–5. A comparison of the species diversity recorded during the 1995 survey with that of the present survey (Table 1) clearly shows a substantial increase in the diversity in most of the dams and rivers since 1995. A decrease in the species diversity during the 2001 survey is only evident for the Sabie River, the Nwaswitsontso River and the Orpen Dam (which forms part of the Nwaswitsontso River drainage system). During 1995, molluscs were found in only eight of the 24 sites surveyed during both years, while mol- lucs were present in all of these habitats in 2001. Taking into account the number of habitats sampled during both surveys, it is evident that B. forskalii was present in 16 of the 24 habitats surveyed in 2001, while it was recovered from only four of the habitats during the 1995 survey (Table 1). Further- more, this snail was recorded from nine of the 18 habitats surveyed for the first time in 2001. With regard to its distribution in 1995, it is clear that it was only found in the southern part of the area, while it was more or less evenly distributed throughout the whole area in 2001 (Fig. 2). In contrast to this, Physa acuta Draparnaud, 1805, which is well known for its invading potential, in spite of exceptionally high mean rainfall for the last five years (Table 4), was present in only two of the habitats, while it was found only once during the 1995 survey (Fig. 3). Aplexa marmorata (Guilding, 1828) which was present only in the Sabie River during the 1995 survey was collected in seven habitats located mainly in the south during the 2001 survey (Fig. 4). Lymnaea columel- la Say, 1817 another invader species, was recorded only three times before 1966 in the Kruger National Park and only once during the 1995 survey (Fig. 5). In contrast to this, it was collected in eight habitats during the 2001 survey. ISSN 0075-6458 3 Koedoe 45/2 (2002) Fig. 2. Distribution of Bulinus forscali in 1995 (O) and 2001 (•). Rainfall monitoring stations (♦). de Kock.qxd 2005/12/09 10:01 Page 3 Koedoe 45/2 (2002) 4 ISSN 0075-6458 Table 1 Habitats of the Kruger National Park surveyed in both 1995 and 2001 Rivers Matjulu Spruit Mbyamiti River Mlambane Spruit Nwanetsi River Nwaswitsontso River Sabie River Timbavati River Engelhard Dam Grid reference 25° 21' 26"S; 31°31'11"E 25°13'27"S; 31°34'38"E 25°21'26"S; 31°31'11"E 24°25"12"S; 31°47'01"E 24°37'10"S; 31°47'28"E 25°07'06"S; 31°55'00"E 24°13'50"S; 31°38'03"E 23°50'28"S; 31°38'07"E 1995 No molluscs Bulinus globosus Spathopsis wahlbergi No molluscs Bulinus globosus Bulinus natalensis Bulinus forskalii Biomphalaria pfeifferi Lymnaea natalensis Burnupia sp Melanoides tuberculata Cleopatra bulimoides Spathopsis wahlbergi Spathopsis petersi Corbicula fluminalis Eupera ferruginea Cleopatra ferruginea Spathopsis petersi Bulinus globosus Biomphalaria pfeifferi Lymnaea natalensis Lymnaea columella Physa acuta Aplexa marmorata Melanoides tuberculata Unio caffer Eupera ferruginea Oxyloma patentissima No molluscs No molluscs 2001 Bulinus globosus Bulinus forskalii Biomphalaria pfeifferi Lymnaea natalensis Lymnaea columella Aplexa marmorata Burnupia caffra Melanoides tuberculata Bulinus globosus Lymnaea columella Aplexa marmorata Bulinus forskalii Biomphalaria pfeifferi Aplexa marmorata Bulinus tropicus Bulinus forskalii Biomphalaria pfeifferi Ceratophallus natalensis Burnupia caffra Melanoides tuberculata Spathopsis petersi Corbicula fluminalis Oxyloma striata Bulinus forskalii Physa acuta Aplexa marmorata Bulinus africanus Lymnaea natalensis Burnupia caffra Bulinus globosus Bulinus tropicus Bulinus forskalii Biomphalaria pfeifferi Physa acuta Melanoides tuberculata Corbicula fluminalis Pisidium pirothi Bulinus tropicus de Kock.qxd 2005/12/09 10:01 Page 4 ISSN 0075-6458 5 Koedoe 45/2 (2002) Table 1 (continued) Rivers Gezantombi Dam Gudzani Dam Klopperfontein Dam Krapkuil Dam Kumana Dam Manzimahle Dam Mazithi Dam Nkaya Pan Nsemani Dam Nwanetsi water hole Olifantdrinkgat Orpen Dam Pioniers Dam Grid reference 25°20'05"S; 31°52'35"E 24°22'05"S; 31°55'33"E 22°37'42"S; 31°10'20"E 23°14'43"S; 31°18'07"E 24°37'11"S; 31°47'40"E 24°53'51"S; 31°43'55"E 24°43'20"S; 31°50'34"E 24°31'54"S; 31°45'25"E 24°23'32"S; 31°43'05"E 24°28'22"S; 31°58'51"E 24°53'56"S; 31°44'54"E 24°47'31"S; 31°53'51"E 23°31'29"S; 31°23'56"E 1995 No molluscs Bulinus globosus Bulinus forskalii Biomphalaria pfeifferi Lymnaea natalensis Spathopsis petersi Unio caffer No molluscs No molluscs No molluscs No molluscs No molluscs No molluscs No molluscs Bulinus globosus Bulinus natalensis Bulinus forskalii Biomphalaria pfeifferi Lymnaea natalensis Burnupia sp. Melanoides tuberculata Cleopatra bulimoides Spathopsis wahlbergi Spathopsis petersi Corbicula fluminalis Eupera ferruginea No molluscs Bulinus globosus Bulinus tropicus Bulinus forskalii Biomphalaria pfeifferi Lymnaea natalensis Ceratophallus natalensis Melanoides tuberculata Spathopsis wahlbergi Spathopsis petersi Bulinus globosus Biomphalaria pfeifferi Lymnaea natalensis 2001 Bulinus forskalii Lymnaea. columella Aplexa marmorata Burnupia caffra Bulinus globosus Bulinus tropicus Biomphalaria pfeifferi Lymnaea natalensis Ceratophallus natalensis Melanoides tuberculata Spathopsis petersi Corbicula fluminalis Bulinus tropicus Bulinus forskalii Burnupia caffra Bulinus forskalii Bulinus forskalii Bulinus forskalii Bulinus forskalii Bulinus forskalii Burnupia caffra Bulinus globosus Bulinus tropicus Bulinus forskalii Biomphalaria pfeifferi Ceratophallus natalensis Burnupia caffra Spathopsis wahlbergi Spathopsis petersi Corbicula fluminalis Bulinus forskalii Lymnaea columella Ceratophallus natalensis Bulinus globosus Bulinus forskalii Biomphalaria pfeifferi de Kock.qxd 2005/12/09 10:01 Page 5 Koedoe 45/2 (2002) 6 ISSN 0075-6458 Table 1 (continued) Rivers Shitlhave Dam Silolweni Dam Vervoer Dam Grid reference 25°08'38"S; 31°19'31"E 24°29'21"S; 31°50'11"E 25°07'07"S; 31°29'29"E 1995 Melanoides tuberculata Spathopsis petersi Corbicula fluminalis No molluscs No molluscs No molluscs 2001 Lymnaea natalensis Melanoides tuberculata Biomphalaria pfeifferi (shell) Bulinus forskalii Burnupia caffra Lymnaea columella Aplexa marmorata Discussion From the results of this investigation it is clear that the high rainfall average recorded for 1995-2001 period had a significant influ- ence on the number of habitats from which molluscs were recorded as compared to the 1995 survey. For the majority of the habitats the mean rainfall for the 2001 period was nearly double the figures recorded for the 1995 period. The decrease in snail bio-diver- sity in the Sabie and Nwaswitsiontso rivers and Orpen Dam (which forms part of the Nwaswitsontso River drainage system, as previously mentioned) could probably be ascribed to heavy floods in these habitats prior to the present survey. In the same localities investigated, B. forskalii was found only four times in 1995 as compared to 16 times in 2001 and was, furthermore, present in nine of the 18 habi- tats surveyed for the first time in 2001. This snail is able to exploit a wide variety of nat- ural and artificial habitats, including the mar- gins of lakes and permanent swamps, but is most abundant in small water bodies and was also collected in alluvial pools associated with rice cultivation in Gambia (Brown 1994). The fact that this snail was found in many more habitats during the 2001 survey may be ascribed to its ability to aestivate through long periods of drought. However, the possibility that some of these habitats were populated by this snail species for the first time due to the high rainfall of the 1995- 2001 period cannot be ruled out. Aplexa marmorata, a confirmed invader species collected only once during the 1995 survey, was present in seven of the localities in 2001, which was investigated during both surveys. Although it was collected several times during the past 15 years in KwaZulu- Natal, it was recorded only once from the Limpopo Province (Dana 1999) at the time of the present investigation. The fact that this species was found in a number of habitats during the 2001 survey seems to suggest that it could be in the process of becoming well established in the southern parts of the Kruger National Park. Whether it has already invaded habitats in the adjacent Swaziland needs further investigation. Although this species is usually reported from water bodies modified by man (Dana 1999), it was present in several rivers and dams in the Kruger National Park In contrast to this, P. acuta, which has a worldwide distribution and which is estab- lished in many African rivers and lakes from South Africa to Morocco, and is considered the second most successful invader snail species in South Africa (De Kock et al. 1989), was recovered from only two habitats during the 2001 survey as compared to the one in 1995. Its distribution pattern in the de Kock.qxd 2005/12/09 10:01 Page 6 ISSN 0075-6458 7 Koedoe 45/2 (2002) Table 2 Molluscs found in waterbodies of the Kruger National Park surveyed for the first time in 2001 Waterbodies Grid reference 2001 Grootvlei Dam 23°20'41"S; 31°30'56"E Spathopsis sp. Jones se Dam 24°51'23"S; 31°45'13"E Bulinus forskalii Ceratophallus natalensis Burnupia caffra Kanniedood Dam 23°08'40"S; 31°27'48"E Bulinus tropicus Bulinus forskalii Lymnaea natalensis Letaba River 23°45'28"S; 31°22'20"E Bulinus forskalii Physa acuta Melanoides tuberculata Mestel Dam 25°06'54"S; 31°12'50"E Bulinus globosus Lymnaea columella Aplexa marmorata Mpondo Dam 25°12'12"S; 31°43'08"E Bulinus forskalii Lymnaea natalensis Lymnaea columella Spathopsis petersi Mtombeni Spruit 24°15'40"S; 31°43'40"E Bulinus forskalii Nshawu Dam 23°31'40"S; 31°29'09"E Ceratophallus natalensis Olifantsdrinkbad 23°19'45"S; 31°19'44"E Bulinus forskalii Piet Grobler Dam 24°13'54"S; 31°38'03"E Bulinus forskalii Rabelais Dam 24°27'30"S; 31°29'58"E Bulinus tropicus Bulinus forskalii Sable Dam 23°56'12"S; 31°14'04"E Bulinus tropicus Bulinus forskalii Shisa Spruit 22°57'32"S; 31°15'01"E Cleopatra ferruginea Melanoides tuberculata Spathopsis petersi Corbicula fluminalis Sirheni Dam 22°56'57"S; 31°14'02"E Bulinus globosus Biomphalaria pfeifferi Lymnaea natalensis Lymnaea columella Lanistes ovum Sunset Dam 25°06'57"S; 31°54'41"E Aplexa marmorata Table 3 Waterbodies of the Kruger National Park surveyed for the first time in 2001 and in which no molluscs were found Waterbodies Grid reference Luvuvhu River 22°26'38"S; 31°13'02"E Ngotso Dam 24°10'31"S; 31°43'09"E Nhlanganzwane Dam 25°14'40"S; 31°58'30"E de Kock.qxd 2005/12/09 10:01 Page 7 Koedoe 45/2 (2002) 8 ISSN 0075-6458 RSA shows concentrations of collections around major urban areas (Brackenbury & Appleton 1993). It is now found in various artificial and natural habitat types and taking into account that this species is not highly specific with regard to its habitat preferences it is quite surprising that it has a relatively limited distribution in the Kruger National Park. However, it occurs commonly in pol- luted water and has therefore been nick- named the sewage snail and seems to have difficulties in establishing itself far from human works or habitats (Brown 1978; Brackenbury & Appleton 1993). This could be one of the reasons for its limited occur- rence in the Kruger National Park, especial- ly with regard to the dams and rivers, which originate in or near the park. Although the drought of 1995 may also serve as an expla- nation for this situation, it must be kept in mind that it had been recovered only three times from the Kruger National Park prior to 1966. Lymnaea columella, which is considered the most successful invader snail species in South Africa (Brown 1967) was recovered from five habitats during the 2001 survey and from only one during the 1995 survey. Scrutiny of its distribution in the whole of the RSA (De Kock et al. 1989) shows clear- ly why it has earned the title as the most suc- cessful invader snail species, because it has successfully established itself in vast areas of the country since its introduction in the early 1940s. Before 1966 it was found in 132 loci, and in 225 additional loci from 1966 up to the present (De Kock et al. 1989). This species was first reported from the Kruger National Park in 1967 from three localities by Oberholzer & Van Eeden (1967) and was collected in only three additional localities during the 2001 survey. In contrast to this, A. marmorata was collected in six additional localities since the 1995 survey. This increase in collection sites over a period of only six years seems to suggest that A. mar- morata could become just as successful as L. columella in establishing itself in this coun- try. Fig. 3. Distribution of Physa acuta in 1995 (O) and 2001 (•). Rainfall monitoring stations (♦). de Kock.qxd 2005/12/09 10:01 Page 8 ISSN 0075-6458 9 Koedoe 45/2 (2002) Fig. 5. Distribution of Lymnea columella in 1995 (O) and 2001 (•). Rainfall monitoring stations (♦). Fig. 4. Distribution of Aplexa marmorata in 1995 (O) and 2001 (•). Rainfall monitoring stations (♦). de Kock.qxd 2005/12/09 10:01 Page 9 Koedoe 45/2 (2002) 10 ISSN 0075-6458 Table 4 Rainfall records of the Kruger National Park at the different monitoring stations nearest to collection sites over different time periods Rainfall monitoring station Berg-en-Dal Crocodile Bridge Houtboschrand King Fisher Spruit Letaba Lower Sabie Mooiplaas Nwanetsi Pafuri Phalaborwa Pretoriuskop Punda Maria Satara Shingwedzi Skukuza Tshokwane Collection site Matjulu Spruit Mlambane Spruit Mpanamana Dam Gezantombi Dam Nhlanganzwanedam Mtombeni Spruit Ngotso Dam Timbavati River Rabelais Dam Engelhard Dam Letaba River Olifants River Lower Sabie Rest Camp Mpondo Dam Sabie River Grootvlei Dam Nshawu Dam Olifantdrinkbad Pioniers Dam Tsende Drif Gudzani Dam Nwanetsi River Nwanetsi water hole Luvuvhu River Sable Dam Mbyamiti River Mestel Dam Shitlhave Dam Klopperfontein Dam Nkaya Pan Nsemani Dam Piet Grobler Dam Kanniedood Dam Krapkuil Dam Sirheni Dam Sisha Spruit Manzimahle Dam Olifantdrinkgat Skukuza fish pond Sunset Dam Vervoer Dam Jones se Dam Kumana Dam Leeupan Mazithi Dam Nwaswitsontso River Orpen Dam Silolweni Dam Average rain- fall of all records avail- able (mm) a 547 386 523 435 577 449 515 332 523 709 528 526 462 533 533 Average rainfall for 1882 - 1983 (mm) a 380 304 265 228 493 196 308 98 234 417 142 326 248 272 295 Average rainfall for 1985 - 1987 (mm) a 521 355 545 335 536 447 437 422 484 428 520 428 356 547 547 Average rainfall for 1991 - 1994 (mm) a 506 279 440 281 441 281 376 297 412 507 327 507 334 411 411 Average rainfall for 1995 - 2001 (mm) 832 814 560 673 418 787 663 653 652 673 989 722 595 704 674 761 de Kock.qxd 2005/12/09 10:01 Page 10 With regard to Lanistes ovum Peters in Troschel, 1845 it needs to be mentioned that large specimens were collected before 1966, while none were found during the 1995 sur- vey. However, juvenile specimens were found in the Sirheni Dam during the 2001 survey which might indicate that this species could be in the process of recovering from the previous droughts. More or less the same conclusions could be drawn for the bivalves on record for the park in general and the Spathopsis spp. in particular. Conclusions From the results of this investigation it is clear that there was a significant increase in the number of localities which yielded mol- luscs during the present study as compared to the 1995 survey. This could mainly be ascribed to the abnormal high rainfall during 1995-2001 period. With regard to the invad- er species A. marmorata, a follow-up survey in about five years’ time is recommended to establish whether the northern parts of the Kruger National Park are also going to be invaded by this species. Acknowledgements The authors wish to thank the following institution and persons for assistance: South African National Parks for permission to conduct the research; Dr Andrew Deacon for coordinating this project on behalf of the Kruger National Park and Mr Nick Zambatis for kindly making the necessary rainfall data available. References BRACKENBURY, T.D. & C.C. APPLETON. 1993. Recolonization of the Umsindusi River, Natal, South Africa, by the invasive gastropod, Physa acuta (Basommatophora, Physidae). Journal of Medical & Applied Malacology 5:39-44. BROWN, D.S. 1978. 35. Freshwater molluscs. Pp. 1155-1180. In: WERGER, M.J.A. (Ed.). Biogeog- raphy and ecology of southern Africa 1. The Hague: Junk. BROWN, D.S. 1994. Freshwater molluscs of Africa and their medical importance. London: Taylor & Francis. DANA, P. 1999. Aplexa marmorata (Guilding, 1828) (Basommatophora: Physidae) — an invasive freshwater snail in South Africa. M.Sc thesis, University of Natal, Durban. DAVIES, B. & J. DAY. 1989. Vanishing waters. Cape Town: University of Cape Town Press. DE KOCK, K.N., P.H. JOUBERT. & S.J. PRETORIUS. 1989. Geographical distribution and habitat preferences of the invader freshwater snail species Lymnaea columella (Mollusca: Gas- tropoda) in South Africa. Onderstepoort Journal of Veterinary Research 56: 271-275. DE KOCK, K.N. & C.T. WOLMARANS. 1989. A re- evaluation of the occurrence of freshwater mol- luscs in the Kruger National Park. Koedoe 41:1-8. OBERHOLZER, G. & J.A. VAN EEDEN. 1967. The fresh- water molluscs of the Kruger National Park. Koedoe 10: 1-42. ZAMBATIS, N. & H.C. BIGGS. Rainfall and tempera- tures during the 1991/92 drought in the Kruger National Park. Koedoe 38: 1-16. ISSN 0075-6458 11 Koedoe 45/2 (2002) de Kock.qxd 2005/12/09 10:01 Page 11 << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJDFFile false /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /ColorConversionStrategy /LeaveColorUnchanged /DoThumbnails false /EmbedAllFonts true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveEPSInfo true /PreserveHalftoneInfo false /PreserveOPIComments false /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputCondition () /PDFXRegistryName (http://www.color.org) /PDFXTrapped /Unknown /Description << /ENU (Use these settings to create PDF documents with higher image resolution for high quality pre-press printing. The PDF documents can be opened with Acrobat and Reader 5.0 and later. These settings require font embedding.) /JPN /FRA /DEU /PTB /DAN /NLD /ESP /SUO /ITA /NOR /SVE >> >> setdistillerparams << /HWResolution [2400 2400] /PageSize [612.000 792.000] >> setpagedevice