Freshwater H'etlands anel their sustainable jillure

Mossakowski, D. (1970). Ökologische Untersuchungen an epigüischen Co CHAPTER 28 atlantischer Moor- und Heidestandorte. ZeitscllrUi/ ~Visse/ls('llajiliche Zool 233-316 Obrtel, R. (1972). Soil surface Coleoptera in a reeel swanlp. Acta Scientiae EFFECT OF FLOODS ON GROUND BEETLES Brno, 6(6): 1-35 Thiele, H. U. (1977). earl/bill Beet/es in their EIH'ironll/cllt. Springer Verlag, H (CARABIDAE) AND fIAI?VESTMEN (OPILIONES) 369 pp. Tietze, F. (1974). Zur Ökologie, Soziologie und Phänologie der Laufkäfer (Co Carabidae) des Grünlandes inl Süden der DDR. V. Zur Phänologie der Cara L. Klinzes untersuchten GrÜnlandes.llercynia, 11: 47-68

Most spccics of carahids and harvestnlen are epigcic and, thus, their würld is only two-dilncnsional. Adults of both groups usually inhabit the surface earth, litter and the lowest layer of vegetation together \vith their prey, i.e. Diptera larvae, gastropods, Collelllbola spp., spiders, isopods and acarids (Thiele, 1977; Adatns, 1984). Life on the soil surface lnay beconle dangerous because of soil surface disturbances, drying or floods. There are severaI kinds of natural events that 1l1ay rcsuh in tClllporal dcstruction orthc epigcic aniJllal enVirOnl11cnt (Galle ef al., 1982). Flooding is an exalnple of such disturbance. In the Wet Meadows area, the environment of epigeic arthropods is overlaid by up to 0.60 m of water during a flood; this milieu is inhospitable and inlpenetrable for n10st epigeic species. Fate 01' the anilnals is variolls. Many species 01' carabids occllrring in flooded habitats are able to survive under the willcr sllrfaec 1'01' two weeks to two 1110nths (Paln1en, 1945, 1949; HeydeJnann, 1967). Some SpeCil11enS ofcarabids are carried away by the water strealTI, species eapable of flying can escape, together with fast running aniInals living not far fronl elevated places. Many specinlens Inay survive on stenlS of tall herbs and on tree trunks for sonle tilne, and will retunl after the flooel has receded. The populations weakcned by floods, or not surviving at all, can bc supported or rc-cstablishcd by anilnals froln thc neighbollrhood not afTected by the flüod. Only a fcw spceies are weil adapted to flooding. Their adaptations are both physiologieal and/ür bchavioural: a diplopod, Gonographis adisi, from thc Alnazon ßasin, survivcs für IIp to 6 nlonths under watcr, respiring the air stored llnder trec bark (Adis, 1986); sonle speeies of beetle survive long periods near the watcr surface (Innler, 1979) 01' even under it for sonle tinle (Thiele, 1977). Oribaticls can also tClnporarily change their 1l1ierohabitat (Bcck, 1969, 1976) and lnany aniInals escape to tree crowns (Adis, 1981, 1982; Erwin, 1983; Friebe and Adis, 1983; Adis and ScheUer, 1984). SOJne Collenlhola spp. survive flood periods as eggs, i.e. at an ontogenetic stage resistant to a nearly anaerobic environnlcnt. Moreover, flooels aet as Clles breaking the egg donnancy in SOlne spccics (Tan1111, 1984a,b, 1(86). In sonle earabids, survival of larvae subjccted to f100ds is llighcr than that 01' adults (Hcydcrnann, 19(7). In grollnd beetles and harvestInen, the two groups treated in this Chapter, both physiological and behaviouraI adaptations can be expected to occur. Ground

360 361 Fres/nvater H'ctlands alld thci,. sllstaillahle.tittllre E.tlcet (~f.t7oods on grolInd beetles and harvestl1zen beetles are 111arked for their wing polYlnorphisnl (Den Boer, 1971; Den Boer et al., 1980). SOlne species are winged and others are not. SOllle species have both winged and winglcss anilllals in their populations. Pterygote specilllens can take advantage of telnporary habitats such as floodcd areas because they are able (I) to escape frorn the area when the water level is rising, and (2) to recolonize rapidly the defaunated arcas when the tlood is over (Roff, 1990). Most harvestmen prefer l11esic and wet habitats, but they only rarely colonize Shrubs flooded areas. The only, so far, known adaptation of opilionids to tlooding is Meadow behavioural; in the tropics sonle epigeic harvestnlcn cscapc to tree crowns for Pond long periods during floods (Friebc and Attis. 19X3). In Ccntral Europe. silnilar Corltour hehaviour has not yct hCCIl ohscrvcd. Tltus. pefllWlll'lll popubliolls ofltarvesIIlH.'1l line are likely 10 bc l'oulld only in habilals \vhich are llo1 affcc1cd by flooding or Streams which are tloodcd only for a short period. If the fauna at a site is either partly 01' cOl1lpletely dal11aged by a tlood, a Fields recolol1ization proccss folIows. Not only different species but also sexes of individual species differ in their ability to spread. According to Fisher's theory (1930) the sex ratio (Illale:fclnale) should be about I: I, because it is beneficial Figure 28.1 Location 01' the sanlpling sites in the Wet Meadows area near TIebon (fuH for outbreeding equilibriuln populations to renlo,:e sIllall deviations fronl this circles); the site of the clin1ate station is indicated by an open circle ratio (Trivers, 1985). This is also true for carabids (Sustek, 1984). In a population which has just been established, one can expect deviations in the sex ratio biased Table 28.1 Basic characteristics of the sanlpling sites situated at different elevations towards I1lales which are nlore 1110bilc and, thus, able to colonize new habitats above 111ean water level in Roznlberk fishpond earlier than females. Populations closc to extinction should bias their sex ratio towards feInales that are nlore tolerant of stress factors (Sustek, 1984). Samp/ing sites It is ahnost ilnpossible to predict the long-tenn dynalnics 01' individual Ahhrel'iatiol1 UD sc AP CG RO populations in the Wet Meadows because or the unpredictable effects of tloods. But it is clear that the water level and probably also other environnlental factors Habitat Rudcral stand Shrub. I1lcadow Mown shorc Wctland Fishpond correlated with it or with soil Inoisture content, tluctuate less widely on drier Elcvation IcmJ 39 IX 16 12 3 sites than near the fishpond shores. This relationship ShOltld be reflected in the Dominant plants Urtica dioica + Sahx pentandra A/opccurus Carcx grad/is Glyccria nUlnber of species with deviated sex ratios. 'Thcreforc, it can be expected that: Sahx cillcrea + Deschampsia pratcllsis + C. l'esicaria maxima (1) the proportion of winged specinlens of canlbicls is greater on tlooded sites, c{)c.\pitosa (2) persisting populations of harvestnlen are found only on sites not subjected to flooding and (3) the proportion of species with deviated sex ratio is greater on frequently flooded sites than in steady-state habitats. winter season. Adults of carabids and both adults and juveniles of harvestmen were included in this study. The nOlnenclature ofcarabid beetles follows Freude elol. (1976), that of harvestnlen Martens (1978). MATERIAL AND METHODS The investigations took place in the Wet Meadows between the town ofli"ebon RESULTS and Rozmberk fishpond (see Chapter 13) froln 1985 to 1986. Five ~cologically differing sites were explored in 1985 and foul' in 1986 (Figure 28.1 and Table The nunlbers of specinlens of carabids and harvestillcil caught on individual 28.1). On each sanlpling site, six pitfall traps of Xenl in dialneter were placed at sites in Wet Meadows arc given in Tablc 28.2. Total nUlnber of carabids, the centre of each site, the other five traps, each 01' 4.5 cm in dialneter, were belonging to 53 species, was 4203. Total number ofharvestmen was 1460; they placed in the corners of a pentagon so that the neighbouring traps were about were represented by 6 species (Table 28.3). The number of captured specimens 2 In apart. The traps were Inade of glass and fillcd up to one third with 6% increased in the second year 01' our investigations in both grollps and on all sites formaldehyde solution. The saIllples were collected lnonthly, except for the exccpt the Ineadow (AP), where the nunlbers of carabids slightly decreased

362 363 Freslnvater l-vetlands and their sustainahle jilture E.tt'ect qjj700ds 011 groulld heet/es lind harvest/11en

Table 28.2 Pitfall trap catches ofcarabid bcclles anel harvestnlen in the Wet Meadows Table28.3 Survcy ofcarabid bcctles and harvestlnen frOI11 pitfall traps and relationship in the period 1985-1986 bctween the nUlnbers of nlacropterous anel brachypterous carabids (nl!b); * significant 2 diffcrencc bctween ycars (chi , p < 0.05). Sec Table 28.1 for abbreviations Sitl' Sill' UD SC AP ce RO UD SC AP CG RO Year 1985 1986 1985 1986 1985 1986 1985 1986 1985 Year 1985 1986 1985 1986 1985 1986 1985 1986 1985 Total Carabidae Agonum jilliginosum 184* 53* 85* 47* 5 77 56* 79* d Carabidac A. moestul1l 14* 2 2 24 36 13 m Spccimcns IIX4 934* 196 317* 413 360 152 334* 313 42()3 A.ohsc"r"l11 85* 31* 3 I d Spl'l'il's .'X .'5 15 10 23 22 10 13 IX 53 Amara COllll1l11l1is 71 (li 4 III Ill/b ().-t 0.5 0.7 3.6* 1.5 104 4.2 4.1 3.1 1.12 27* m Bemhidion assimile °l>iliones 152* 144* 29 151* 112* 140* 10 136* 41 d Carahus granulatus Adults 162 570* 24 7* 6 11* I1 69* 2 862 44 3 84 30 2 2 b Dyschirills gioboslis I Juveniles 199 231 13 42* 9* 38 66* 598 Elaphrus ClipreliS 3 I 48 m Species 5 5 3 3 4 5 3 3 2 6 Loricera pilicornis 9 18 14 33 48 6 3 m Oodes helopioides 6 15 1 8 2 I 15 35 19 m Poecilus \'ersicolor 5* 8 14* 35* III Pterostidllls diligells 93 30 6 12 4 2 14* 9 d P.minor I 2 9* 10 12 35* 32 l1l crabk 2X.3). Thc incrcasing trend 01' capturcd Spccilllcns on thc sites conlpared P.nigrita 2 8* 39 51* 13 12 7 9 19 m followed the elevation gradient in both arthropod groups studied (RO < SC, P. strenll"s 62* 59 19 41* d P. me/anarius 75 96* 47* 6 d CG, AP « UD). Nevertheless, elevation nlay not be the only factor affecting Trechlls secalis 328 333* 2 I1 10 5 b the nUlnbers; the other factors Inay be the distance froHl Rozmberk fishpond Opiliones (adults) shoreline, presence 01" shrub cover, and regularity or 111anagelllent of the wet Lacinills ephippiatlls 39* 38 13 2 1 I 4 29* grassland. Species richness was correlated with the numbers of specimens in Nemastoma 11Igubre 33 225* 3 I both groups, except for markedly high nunlbers of carabid species captured in Opigolophus tridens 89* 245* 9 2 I 1 7 14 Rilaena triangularis 62 6 2 26* the nleadow (AP) in comparison with the other two sites situated at similar ... elevation (SC and CG) - see Table 28.3 and Figure 28.2. * sex ratio (males/females) differs from I: 1(chF, p< n.05) Thc rclationship bet wccn thc nUlnbcrs or 111acroptcrous and brachyptcrous - UD, SC, AP, ca, RO - sec Table 28.1 - m - macropterous species, b - braehypterous, d - dimorphie species of carabids specilllens ofcarabids differed between the habitats studied (1985: chi2= 211.6, _. presenee ofcarabid specics not reaching thc dominancc 01' 5% in any stand: Agollllm gracile (CG, DF= 4, p< 0.0005; 1986: chi2= 197.0, DF= 3, p< 0.0005): brachyptcrous UD - * in 1985); A. liven.\' (SC * in 1985, UD, CG); A. miclIns (UD, SC); A. picellm (RO*); speciInens prevailed on the driest site (UD) in both years. High proportion of Amara anthobia (UD); A. aulica (UD); A. cllrta (AP); A. Ilinicollis (UD, AP); A. ovata (UD); A. similata (UD, SC, AP); Bembidion glIttlIla (UD, AP); B. IWllIlatlim (UD, AP); B. O!JtllSllJJl (UD, Inacroptcrous Spccilllcns was found on each of the three sites situated in the AP); B. lInicolor (UD); Blethisa mliltipunctalll (RO); Ca/at/ws mclalloceplwills (UD, AP); Chlaenitls area close to the fishpond. These nUIllbers did not change significantly between nigricornis (UD, AP); CIi\'illa ('olltracta (UD, AP); C. fossor (UD, SC, AP, RO); Demetrias 1985 and 1986 except for Sahr scrub (SC) where the proportion ofl11acropterous mOl1ostigma (UD); Dicheirotrichlls rll.JitllOrax (UD, Ca); Dromills sigma (UD); Harpaills aelleus (AP); Leistus rufescells (UD); L. sp. (UD); Patrohlls atrorujils (UD); Poeci/lls clipreus (UD, AP); individuals decreased. Pterostichus anthracillus (UD, AP); P. gracilis (UD); P. Iliger (UD, CG); P. ob/ongoplIllctatus The species of carabids split into two grollps with different timing of their (UD); P. l'erllalis (UD, AP); SYllucllllS nh'alis (UD); Treclllls qlladri:ariatlls (UD) ontogenetic cycles: nlost species present on the Wet Meadows sites over-winter - rare opilionids: Paranemastoma qlladripllnctata (UD); Plwlallgiul1l opilio (UD, AP, RO) as larvae and breed in autuIlln. A few species of the other type, i.e. hibernating as adults and breeding early in the season, were found only on the driest site (UD). However, since the species nUlnbers were rather low, the difference between those two groups was non-significant. If the comparison were based on individuals instead of species the proportion 01' autumnal breeders on individual sites would beconle Inarkedly different. Alnong harvestnlen, the species also differ in their life history. Rilaena Iriangu/aris hatches in late SUlllJ1ler and the aninlaJs Illature during the next

364 365 ~vetlallds Fres/nvater and their slistainahle .tiaure E.lI(Jct (~t'.t70ods 0/1 grolllld beetles alld harveSII71en

Lacinius ephippiatus 1985 1986 §~~ 8 ~ 30 \ \tl/ o z o CJ)

J FMAMJ J ASONDJ FMAMJ J ASOND Months

o Rilaena triangularis z 1985 1986 o CJ)

o ol.-.y--.--.....-.....-.---.--...... ,.....-.---.-...... -+--.-...... -.--..,...... ,...--.---.--.--.....-.---.---. z JFM AM JJA so ND J FM AM J JAS 0 N D o Months CJ)

spring; no adult specil11en of this species was caught in ~ 985, in contrast to high nun1bers collected in the subsequent year (Figure 28.3). Two other abundant species, Lacinius ephippiatus and Oligolophus tridens, are summer and autumnal breeders, respectively; el11ergence of their adults was delayed by about one 000009 ~ ~ ~ Cf) ~ r-4 ~ .. ~ g 0 n10nth in 1985 as con1pared with 1986. Both nenlastoillatids and Phalangiunz ~JltleJadtual ue~w [Ja] [ww] UOHl.!JP!JaJd opilio were too rare for any delailed exanlinalion. The sex ratio devialed [rol11 that expected (1: I) in 11 out of 16 comnlon carabids and in all four COlnnlon harvestnlen. This ratio, however, did not depend on site (Table 28.4). There \vere only two species of carabids with variable sex

366 367 Fres/nvater };vetlands and their slistaillahle.flaure E.JfeCI offloods on gral/nd beetles alld harveslll1en

Table 28.4 Sex ratios (rnales/fernales) in C0011110n carabids and harvestnlen collected COMPARISON WITH OTHER DATA with pitfall traps (1985-1986); the effect of salnpl ing sites on sex ratio was tested by 2 Carabids hClcrogcncily (H-chi ), :I: - P < 0.05; n.s. non significanl Species assel11blages of carabids and harvcstnlcn have been studied in ITIany l1ulle ./{,l1ulle P II-ehF! DF P habitats by nUlnerous authors, since it is relatively easy to collect a lot of Carabidae SpeCil11CnS with liHle effort. Nevertheless, wetlands are insufficiently represented Ag0l1U111 jiiliginosull1 1:2.16 * alllong the habitats studied, because the standard 111ethods ofinvestigation often A. 1110estunl 1: 1.47 n.s. 1.69 2 n.s fail here, whel1 catches are s0I11etillles washed away by fIoods. This was also 4.80:1 * A. ohscurul1z the reason why the RO site was not accessible fr0l11 the end of 1985. AI11ara COluI11unis I: 1.1 () n.s. BClllhit/ioll (/ssi/llile 2.X(): I :1: Thc silllilarity ill specics corllpositioll 01' carabid assclnblagcs in non-forestcd Carabus gralllllatlis 2.41: 1 * 12.38 4 * wetlands is great in Central Ellrope (conlpare Stiller, 1932; Avasi, 1987; Chapter Elaphrlls ('uprclis 1.56:1 n.s. 27). Thc specics richncss is ralher low in the Wet Meadows area. A few weiland I: 1.17 n.s. 0.16 2 n.s Loricera pilicorllis species, represenled by Godes helapioides, AgOllU111 subgen. Europhilus, A. Oodes helopioides 1.54: 1 0.29 3 n.s * livens, Elaphrus cupreus and sOlne species with shortened elythrons, were Poecilus eaerulesee/ls 1:2.81 :/: 0.08 I n.s Pterostichus dilige/ls 1: 1.70 :1: 2.53 4 n.s accolllpanied by I11eadow species, the abundance of which Inarkedly increased P.111inor ]: 1.46 * 20.56 3 * in the season without flooding (1986) and was high on the driest site (UD). P.nigrita 1: 1.35 n.s. 10.19 4 * High proportions 01' brachypterolls anilnals have been reported fronl: P. stre/lliliS I: 1.87 :/: 0.11 1 n.s. (I) woodlands not associated with water, (2) forested swanlps and borders of 3.23:1 2.63 1 n.s. P. Inelanarius * lakes within forests, (3) under logs, stones and burrows, (4) in caves (but not in Treehus seealis 1.33: 1 1.81 1 n.s. * ant nests), and (5) in thc alpine zone and tundra (Roff, 1990). Prevalence of Opiliones (adults) nlacropterous carabids is expected in any unpredictable or unstable habitat. 11' Laeinius ephippiatus 1:2.81 :-i: 0.64 2 n.s. thc habitat becoIlles Illore stable, then the nlacropterous animals are step-by­ Nel11astonla lugubre 1:2.86 * Opigolophus tridens 1:3.41 * 5.9 2 n.s. step rcplaccd by brachypterous ones that save energy by not producing wings Rilaena triangularis I: 1.91 * 17.51 2 ~: with energy-deIllanding nluscles and thus, their reproductive effort can be higher than in nlacropterous anilllals (Roff, 1986, 1990). In lhe Wct Meadows, which have becolne SOIllcwhat drier during the last three decades of the twentieth century, brachypterous anilnals are probably ratios deviating from 1: 1, Carabus granulatlls with a Inale-biased ratio and expanding and colonizing new sites, replacing the original residents. Pterostichus n11l1or with a fenlale-biased ratio. In the fOrIller species, the highest Nevertheless, floods can, froIll tirlle to tirne, inteITupt 01' even reverse this process. ratio was found in Carex wetland (4.84: I) and thc lowest \vas on the fishpond Accürding to M liller (1970) and Sustek (19X4), the sex ratio is a vaillable shore (1.56: 1). In the latter species, lhe sex ratio was very variable, belween indicator 01' state and dynaillics 01' a carabid population. Thc species are expected l11ale-biased (RO - 1.91: I) and fClnalc-biascd (SC - I:2.XO and CG 1:2.92). lo adjust to deviations in the sex ratio (Fisher, 1930). Male-biased sex ralios The nunlber 01' species wilh devialed sex ratios on individual sites was highesl were, however, found in expanding populations and feIllale-biased ratios were on the driest site (UD) but was also correlated wilh the nUlllber 01' captured in relreatin,g ones (Sustek, 1984). In our locality thc sex ratio cleviales specilnens and thus, the power 01' lhe chi~ test differed betweell siles. As lhe signiricantly frol11 thal expecled (1: 1) in lwo lhirds 01' the carabid species (Table number of COlTIlTIOn species both with the expected and deviated sex ratios was 28.4). FeIllaies prevail in six species, Illales in five. Our findings show no trend nearly the sanle on each site, there is hardly a trend 01' one-sex-deviation towards of those biases either within or bctwcen the sites (coll1pare Sustck, 1984 für the driest site. different conclllsions). Many reasons can bc found for deviations in sex ratios In common harvestmen, the sex ratio was felllale-biased. The relationship in our area. In outbreeding populations, deviations can be caused by differential did not depend on the habitat except for Rilaena triangularis with an equilibrium expense 01' different Illortality of the two sexes. The-latter is assumed to operate (Sustek~ ratio on the driest site (UD - 1.03: 1) and felnalc-biascd ratio on lhe other sitcs in carabids 1984). Another cause nlight be differential lTIigration rate (SC - 1:5, ca - 1:25). of the sexes, since lhe frcquency of winged individuals is usually higher ill Inales (Roff, 1990).

368 369 f"'resll'water wetlands alld their slIstaillohlc .titIlire t..lj'eCI off/oods 0/7 groulld beelles alld harvestnzen

Different factors and processes affecting sex ratios seem to operate in Adis, J. (1986). An Haquatic" Inillipede frolll a Central Anlazonian inundation forest. individual species and detailed population studies are required to explain the Occologia, 68: 347-9 observed pattern cOlnpletely, both in the sex ratio and in the proportion of Adis, J. and Scheller, V. (1984). On the natural history anel ecology ofHanseniella arborea brachypterous carabids. We can speculate that the differential 1110bility and Scheller (Myriapoda, Symphyla, Sculligerellidae), a 111igrating symphylan from an 1110rtality ofsexes result in changes in population structure within shorter periods Alnazonian black-watcr inundation forest. Pedobiologia, 27: 35-41 Andersen, J. (1985). Hunliclity responses and water balance of riparian species of than do the benefits or retreats caused by either nlacroptery 01' brachyptery. Belnbidiini (Col., Carabidae). Ecol. EIl/0J1l01., 10: 363-75 Intennittently flooded telnperate wetlands are Inostly inhabited by spring Avasi, Z. (1987). Flood as ecological perturbation of epigeic animal communities. 11. 1939) breeders of carabid (Larsson, because only they are able to finish their The effect of flood on ground beetle asselnblages (Coleoptera: Carabidae). Tiscia, developlnent within the period between the winter and SUlllnler floods (Andersen, 22:99-107 1985). In Ollr study area. the selcctioll operating agaillst alltlllllllal breeders seelns HeCK. L. ( 19()1..). Zum jahreszeitlichen Masscnwcchsel zweicr Oribalidenarlen (Acari) to be quite strang on all sites except for the driesl (U[)). Silllilar reslllts \verc im lIeolropischcll Üherschwcllllllullgswald. VerhalldlungcJI der !)l'ulsc!teJl obtained by Pizl (Chapter 27), Obrtel (1972) and Krogerus (194g). Loologiscl1cJI G'cscl/schaji, 1968: 535-40 Beck, L. (1976). Zlun Massenwechsel zweier Oribatidenarten (Acari) im neotropischen Überschwenlnlungswald. AJ11aZoniana, 6: 1-20 Harvestnlen dcn Bocr, P.l. (1971). On the dispersal power of carabid beetles and its possible Sex ratio in harvestlnen caught by pitfall traps is usually felnale-biased except signiticance. In den Boer. P.J. (cd.), Dispersal and dispersal power ofcarabid beetles. for species with very long legs whcre nlales prevail since their Inobility is Nlisc. Pup., 8: 119-135 den Boer, P.J., van Huizen, T. H. P., den Boer-Daanje, W., Aukema, B. and den Bieman, nlarkedly higher than that offemales (e.g. Martens, 1978; KI ilnes and Spicakova, C. F. M. (1980). Wing polynlorphisln and dilnorphisln in ground beetles as stages in 1984). High variability ofthe sex ratio in between the sites Rilaella triollgularis an evolutionary process (Coleoptera, Carabidae). EnlOmolof!Ja generalis, 6: 107-34 01' is striking. In wetlands (CG) nlales were nearly abscnt. I-larvestlnen without Erwin. T.L. (19X3). Beetles anel other insects of lropical forest canopies at Manaus, alnl0st without nlales are rare but sonlctinlcs persistent since egg fcrtilization is Brazil, by inseclicidal fogging. In Sutton, S.L., Whiunorc, T.C. and Chadwick, A.C. not required in those populations (see Martens, 1978 and references therein). A (eds), Tropical Rain Forest: Ecology and Managel1lent. Blackwell, Oxford, pp. 59­ different explanation tnay be proposed for the Inarkedly fenlale-biased sex ratio 75 in Rilaena triangularis. The season of 1984 was without any strong and Fisher, R.A. (1930). The Gellctic Theory ofNatural Selection. Clarendon Press, Oxford prolonged flood. One year later, in spring 1985, two severe floods occulTed, Freude, H., Harde, K.W. and Lohse, G.A. (1976). Die Käfer Mitteleuropas. Band 2. inten-upted by a dry period with elevated tenlperatures. Nevertheless, that year Adephaga I. Goedke and Evers Verlag, Krefeld no adult specimens of R. triangularis were caught in the Wet Meadows. In Friebe, B. and Adis, J. (1983). Entwicklungszyklen von Opiliones (Arachnida) inl autumn 1985, 210 juveniles of that species were collected and in 1986 adults Schwarzwasser Überschwelnnlungswald (lgapo) des Rio Taruma Mirim (Zentralanlazonien, Brasilien). 8: 0 were found on all sites. This suggests that in March 1985, juveniles of Al11azoniana, 10 I-I Galle, L., Györffy, G. and Hornung, E. (1982). Flood as an ecological perturbation of R. triangularis left their winter fefuges and were then deciInated by the long epigeic aninlal cOlnnlunities. 1. SOlne preliIninary hypotheses on the application of and repeated floods. SOlne individuals ofR. triangularis escaped or recolonized catastrophe theory by evaluating some Matelz-Körtvelyes data. Tiscia, 17: 155-62 the area in SUlnmer and thus, in aututnn nUlnerous hatched juveniles were caught. Heyclenlann, B. (1967). Die biologische Grenze Land-Meer inl Bereich der Salzwiesen. Next year, the developlnent of the R. triangularis population was not disturbed Steiner, Wiesbaden since only a short flood came in lune, and the speciInens of Rilaena were able lnnler, U. (1979). Abundance fluctuations on habitat changes of soil beetles in Central to leave the earth surface and survive the flood on vegetation (see Klilnes, 1990 Anlazonian inundalion forests (Co!.: Carabidae, Staphylinidae). Studies on Neo/ropical for details). Faulla alld Enl'ironnlent, 14: 1-16 Klilnes, L. (1990). lnlpact of floods on the life cycle in Rilaena triangularis (Herbst) (Opiliones). Shorllfk Jihoceskeho Muzea v Ceskych BudejovicEch, PfErodnEVedy, 30: REFERENCES 37-45 (In Czech with English sumnlary) Adatns, J. (1984). The habitat and feeding ecology 01' woodland harvestlnen (Opiliones) Klilnes, L. and Spicakovä, E. ( 1984). Beitrag zllIn Erkennen der Dynamik von Arachno­ in England. Oikos, 42: 361-70 fauna an denl Waldökoton. Acta Universita/is Palackianae Olol1l11censis Facuftas Adis, J. (1981). Comparative ecological studies of the terrestrial arthropod fauna in RenlJ11 NaturaliuJ1l, Biologica, 81: 167-90 (In Czech with Gennan summary) Central Aluazonian Inundation. AJnazoniana, 7: 87-t73 Krogerus, 1-1. (1948). Ökologische Untersuchungen über Uferinsekten. Acta Zoologica Adis, J. (1982). Eco-entomological observations fronl the Anlazon: 11 Canlbicls are adapted FCllnica, 53: 1-157 to inundation-forests. The Co!llopterist's Bulletin, 36: 440-I

370 371 Fres/Ht'a/er H'etlands and tIJeir slI.staillahle .Iil/ure

Larsson, S.G. (1939). Entwicklllngstypen und Entwicklungszeiten der Dänischen CHAPTER 29 Carabiden. Entonlologiske Meddelelser, 20: 277-560 Martcns,1. (1978). Weberknechte, Opiliones. In Dahl, F. (cd.), Die Tierwelt Deutschlands 64, Gustav Fisher Verlag, Jena, pp. 1-464 (AI~ACHNIDA) Müller, G. (1970). Der Sexual-Index bei Carabiden als ökologisches Kriteriul11. COMMUNITIES OF SPIDERS

Entolllologische Berichte, 1970: 12 <- IN THE WET MEADOWS Obrtel, R. (1972). Soil surface Coleoptera in a reed swarnp. Acta Scielltiae Naturalis Brno,6(6): 1-35 Palmen, E. (1945). Über Qllartierwechsel und sublnerse Über-Winterung einioer V Rüzicka terrestrischer Uferkäfer. A/ll1ales E/lt()/110Iogici Fennici, 11: 22-34 b Pahnen, E. (1949). Fclduntcrsllchungcn lind Expcrilllente zur Kenntnis der Übcrwinterun1!.

cinip.er urerarthropodl'll. i\1I1101t'S 1'.'IIIOIIIO/og;('; /'('1111;(';

Locality Mol - Molillia-doluinated grassland no. I An unlnanaged nloist nleadow with scattered solitary willows (mostly Safix cinerea) and Seots pine (Pinus sylveslris). It is situated in .the castern part 01' the inundation area 01' Rozlnberk fishpond. This 111cadow Inay be classified as a cOlnlllunity of the alliance Molillion coeruleae. Plant species typical of this conllllunity are Molinia coerulea, Sanguisorha o/fici/ut/is, Achillea ptannica, Pellcedanl/lll pall/stre, Juncu.s jiliforlllis, Agrosti.\' canina, Deschanlpsia caespitosa, Senecio ril'ldaris, flydrocotyle vulgaris, Iris pseudacorus, Juncus e.tfilsus, Lysilnachia \'ldgaris, Polen/illa erecta and N(llllllburgia thyrsiflora.

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Copyright 2002 i.!) UNESCO The designations employed and the presentation of the material throughout this publication do not imply the expression of any opinion whatsoever on the part of the pllblishers and the UNESCO Secretariat conccrning the legal status of any cOllntry 01' territory. city 01' area 01' 01' its CONTENTS v authorities, the delimitations 01' its frontiers or boundaries. The authors are responsible for the choice and the prcsentation of the facts contained in this book PREFACE ix anel for the opinions cxprcsscd therein. which are not nccessarily those of UNESCO and do not COllllllit the organizution. EDrr()I{S' PI{EFACE xiii Mention ora trademark or a proprietary product docs not constitute a gllarantee or a warranty or the product and does not imply its approval to the exclusion of other products that also may be suitable. DEDICATION xv No part 01' this publication may be rcproduced. in any form. FOI{EWOI{D xvii without permi.ssion from the publishers except for the quotation or brief passages 1'01' the purposc 01' review. J. Kvet, J. Jenfk and L. Soukupova LIS'T OF CONrrl{IßUT()RS xxiii

ßritish Library Cataloguing in Publication Data Section 1 The key role of \vetlands in the biosphere reserve Fresh water wetlands and their sustainable future: evidence from the Trebon Basin Biosphere Reserve. - (Man arid the biosphere series~ v. 28) I. Trebon Biosphere Reserve 2. Wetland ccology Czcchoslovakia Casc studies 3. Sustainable development Czechoslovakia Case studics 1. HUMAN IMPACTS ON TUE TREBON BASIN BIOSPHERE 1. Kvct, J. (lan). 19lJ3-11. Jenik, J. Uan) RESERVE 3 333.9' 18'09437 J. Jenfk and J. K\'cl ISBN 1-85070-550-X 2. WETLANDS OF THE TREBON BASIN BIOSPHERE RESERVE IN TUE CENTRAL EUROPEAN CON1~EXT 11 J. Jenfk, R. Kurka and S. Huscik

Libnll"Y uf Congrcss Cat&lluging-in-Puhlicatiul1 Data 3. DEVELOPMENrr OF IHP AND MAB STUDIES IN TUE Frcsh water wetlands and their sustainablc future: evidence from the Trebon Basin rrltEH()N BASIN BI()SPHEI{E I{ESEI{VE 19 Biosphere Reserve I edited by Jan Kvet. Jan Jenik. J. KvcI and J. Jen!/.: p. cm. - (Man and the biospherc scries) ISBN 1-85070-550-x 1. Wetland conservation - Czcch Republic - Chranena krajinnä oblast Teeboesko. 2. Section 2 Fishpond luanagement: ecological consequences Wetland ecology Czech RCPllblic - Chrancna krajinna oblast Teeboesko. I. Kvet, J. (lan), 1993- 11. Jcnik, Jan. IU. Serics. 4. UYDROBIOLOGICAL EVALUXfION OF TREBON QH 77.C94 F74 1999 577.68'09437 1--<.Ic2 I 9lJ-046240 FISHPONDS SINCE THE END OF THE NINETEENTH CEN1~URY 31 L. Pechar, I. Pfikryl and R. Faina Typesct by H&H Graphics. Blackbllrn, UK Prinled and bound by Bookcraft (Balh) Ltd., Midsomer Norton, UK v