G. Straatsma*, S. Egli**, 1. KrisaiGrei1huber*** & J. Baar* Applied Plant Research, Mushroom Research Unit, Horst, the Netherlands **
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1 NATURAL RESOURCES OF EDIBLE AND OTHER USEFUL MUSHROOMS IN FORESTS IN RELATION TO PRODUCTIVITY AND BIODIVERSITY G Straatsma*, S Egli**, 1 KrisaiGrei1huber*** & J Baar* * Applied Plant Researh, Mushroom Researh Unit, Horst, the Netherlands ** Swiss Federal Researh Institute WSL, Birmensdorf, Switzerland *** Institute of Botany, Vienna, Austria Abstrat: The variation in the produtivity of speies over the years in a Swiss forest plot and in 13 Austrian forest and grassiand plots were analysed in relation to the overall number of fruit bodies and to the number of speies of the plots The produtivity of most speies in a ertain year were related with the speies rihness of that same year We onlude that speies rihness is a very usefull parameter to express produtivity in general terms The relation between the overall number of fruit bodies and the number of speies seems to represent a ommon relation in ombinatorial mathematis The aumulation of speies rihness in time was modelled The model implies that speies rihness will ontinue to inerease in time The produtivity of speies was haraterized in terms of abundane and yearly frequeny Many speies showed preferenes for speifi years apart from the rihness ofthose years Keywords: abundane, yearly frequeny, range, speies rihness, rarity Introdution Several valuable edible fungi, in partiular etomyorrhizal speies like Boletus edulis and Cantharellus ibarius, an not yet be produed agronomially Knowledge of their produtivity in relation to environmental onditions is of strategie value for the development of (semi-natural) prodution systems (Hall et al 2003) One aspet is the variation in produtivity of a speies in a site from one year to another (alled flutuation by Arnolds 1 992) We were interested in the yearly variation of individual speies and their relation to the variation of overall produtivity, as the number of fruit bodies, and as the number of speies of the mushroom assemblage of a site A data set on one Swiss plot and a data set on 1 3 Austrian plots were made available by Simon Egli and Irmgard Krisai-Greilhuber, indiated as Swiss and Austrian data, respetively Fasinating relations between parameters alulated from the primary data were found, implying that the data sets were highly strutured This would allow for the mathematial modelling of aspets of produtivity and of speies rihness We elaborated this for the aumulation of speies rihness in time as a new approah to quantify the biodiversity of mushrooms of sites Available data Swiss data One forest plot was surveyed for 2 1 years A total of fruit bodies were ounted, representing a total number of408 speies The data were provided by Simon Egli Austrian data Seven forest and six grassland plots were surveyed for seven years The numbers of fruit bodies were estirnated, the number of speies among plots varied from 83 to 292 speies, the total number for all plots was 886 speies The data were provided by Irmgard Krisai Greilhuber 80
2 24 The analyses of the data sets have been desribed separately by Straatsrna, Ayer & Egli (2001) and by Straatsrna & Krisai-Greilhiiber (2003) Years; relation between the number of fruit bodies and the number of speies In the Swiss plot, the total number of fruit bodies and the total number of speies over the years ar orrelated In a log/log plot the relation seems linear (Fig 1),J 5O lowest 182 fr-bod, 18 sp highest 8467 fr-bod, 179 sp 22 Iog logplotted ID ) E 50 s d00 4d00 6d00 8d abundane, number of fruit bodies 12 Fig 1 The relationship between the total number of speies and the total number of fruit bodies in the Swiss plot, over 21 years The same data an also be expressed in aumulative form (the seond dot showing the number of fruit bodies and the number of speies of the first and seond years ombined, et) Again an almost linear pattern appears in a log/log plot (Fig 2) 1 1 : lstyrl89lfr-bod,8lsp fr-bod,408sp/ 21 2 / (_) 0 / // / A umulative abundane, log Fig 2 The relationship between the total number of speies and the total number of fruit bodies in aurnulative form in the Swiss plot, over 21 years 81
3 Fig 4 Randornization of years (using EstirnateS, Colwell 2000) provides for a goöd model of atual aurnulation (Swiss data set) 20 A--randomization years 21 = fruit bodies This number an be used to extrapolating Fig 2 The result from the the development of aumulated speies rihness in time (Colwell 2000, Gotelli & Colwell In a log/log plot the number of speies ofwords and the total number ofwords are (almost) linearly related Fig 3 The struture of a ombinatorial system (Changizi 2001) for whih we analyzed an etext as an example etext Projet Gutenberg 59 sentenes 4arx&Enge1s F words of2216 speies speies ofwords (ommon speies: the, of, in, and, a et; Fig 3) found this pattern to be ommon in ombinatorial systems in whih omponents (fruit bodies) ar ombined to instantiate expressions (speies) For omparison we analyzed an eletroni text, downloaded frorn Projet Gutenberg s website, and we ategorized words into Suh linear patterns ar known from other fields of study as weil Changizi (2001) Obviously an aumulation urve starts with one fruit body of one speies thus in a 2001) In the Swiss survey were found in 21 years On average a year shows log/log plot the urve should start in the origin The aumulation urve an be used to predit randomizations ofyears (Colwell 2000) fits weh to the atual aumulation data (Fig 4) a umulatednunibers - () O ld CID sample based aumulation - ---atua1 aumulation urve 1 V 82 abundane, log / number ofwords in a sentene, log
4 _ Speies; abundane, yearly frequeny and range of distribution Gaston & Blakburn (2000) provided evidene that loal abundane of speies is related with speies range for different kind of speies assembiages Böth in the Swiss survey as weil as in the Austrian survey a strong orrelation exists between abundane and yearly frequeny of speies (Fig 5) o n 4 : // yearly frequeny 2 0 ::Z Fig 5 The relationship between the produtivity of speies, as number of fruit bodies, and their yearly frequeny (maximum number shown in any year, averaged over speies with the same yearly frequeny) [left: Swiss plot, 21 years; upper right 7 Austrian plots in Wiener Wald, lower right 6 Austrian plots in Danube flood plain, 7 years] This makes yearly frequeny an interesting parameter to express (loal) abundane Yearly frequeny is quantified in any survey spanning several years independent from atual abundane ounts 1 estimates The yearly frequeny of speies in the Austrian survey orrelates very weil with their presene in the number of plots being studied Table 1 Distribution of speies over ranges frorn rare to produtive (presene in plots and years; 13 Austrian plots over 7 years) yearly frequeny maximum number ofplots numbers ofspeies total total
5 84 LL - 5 yearly frequeny o I: \ JJ \ 1 1 speies are ommon (Fig 6) Many speies have bw abundanes (yearly frequeny) and are rare, only a few j 0 : express their maximum produtivity then (Fig 7) in the Swiss study shows 194 speies of a total of 408 Even in poor years sorne speies number of fruit bodies in that year Thus years differ in produtivity and on average speies prefer the rihest year However many speies do follow this rule The most produtive year Fig 1 shows that the number of speies within in a year is related with the total Interation between years and speies 05 NN \\ Fig 6 The number of speies within eah yearly frequeny dass [left: Swiss plot, 21 years; upper right 7 Austrian plots in Wiener Wald, lower right 6 Austrian plots in Danube flood plain, 7 years] : I (_) Fig 7 The number of speies showing their maximum produtivity versus the produtivity of the yeais (taken as number of speies) years produtivity, nr speies (log) o see if indeed their range is related with loal abundane(s) lt would be very interesting to study large sale distribution of mushroom speies to
6 Produtivity of groups of speies of different nutritional strategy The Swiss data show quite learly that the two groups of rnyorrhizal and of saprotrophi speies flutuate in onert in produtivity over the years (Fig 8) The Austrian data show the same pattern 25 rnyolthizal saprotrophi : \/ \ / year number nr saprotrophes, log Fig 8 The numbers of rnyorrhizal and saprotrophi speies are related over the years The two groups have different nutritional strategies and the relation found indiates that the variation in produtivity of both groups is/are governed by the same fator(s) and that nutritional strategy and flutuations in amounts of substrate ar not very important in this respet Conlusions We feel that a mathematial approah of biodiversity (Magurran 1988, Rosenzweig 1995, Colwell 2000, Gaston & Blakburn 2000) of fungal assembiages is important for a better understanding of the produtivity of speies of interest and of the rarity of so many speies We onlude that numerial data on fungal assembiages ar highly strutured (Figs 1, 2, 5, 6, 7) Rarity is probably a ombination of bw loal abundanes and a srnall range (Table 1 in ombination with Fig 5) Differenes in produtivity of total numbers of fruit bodies and of numbers of speies were observed over the years (Fig s 1 and 8) On average produtivity of speies and of years ar related; however, rnany speies prefer years apart frorn their general produtivity (Fig 7) Referenes Arnolds E, 1992 The analysis and lassifiation of fungal ommunities with speial referene to marofungi In: W Winterhoff (d), Fungi in vegetation siene Kluwer, 7-47 Changizi MA, 2001 Universal saling laws for hierarhial omplexity in languages, organisrns, behaviors and other ombinatorial systems Journal of Theoretial Biology, 211:
7 86 Projet Magurran AE, 1988 Eologial Diversity and its Measurement Croom Helm 1 Prineton from samples Version 6Obl User s Guide and appliation published at: Colwell RK, 2000 EstimateS: Statistial estirnation of speies rihness and shared speies Gaston KJ, Blakburn TM, 2000 Pattern and Proess in Maroeology Blakwell Gotelli NJ, Colwell RK, 2001 Hall IR, Yun W, Amiui A, 2003 Cultivation of edible etomyorrhizal mushrooms Trends in Biotehnology, 21: and distribution of fungal fruit bodies in a seven year plot based survey near Vienna 53(3): 35 Rosenzweig biodiversity: proedures and pitfalls in the measurement and omparison of speies rihness Eology Letters, 4: Gutenberg ML, 1995 Speies Diversity in Spae and Time Cambridge Straatsma G, Ayer F, Egli S, 2001 Speies rihness, abundane, and phenology of fungal fruit bodies over 21 years in a Swiss forest plot Myologial Researh, 105: Straatsma G, Egli S, Ayer F, 2001b Wie Pfifferlinge im Wald gedeihen Unser Wald, Straatsma G, Krisai-Greilhuber 1, 2003 Assemblage struture, speies rihness, abundane, Myologial Researh, 1 07 : Vogt KA, Bloomfield J, Ammirati JF, Ammirati SR, 1992 Sporoarp prodution by with emphasis on forest eosysterns In: GC Carroll & DT Wiklow (eds), The fungal ommunity Blakwell, basidiomyetes, Quantifying
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