Pseudoscorpions (Arachnida: Pseudoscorpiones) in Strict Forest Reserves in Hesse (Germany)

2015·10.5 MB·English

by  Christoph Muster

5
Arachnologische Mitteilungen 50: 37-50 Karlsruhe, November201
Pseudoscorpions (Arachnida: Pseudoscorpiones) in Strict Forest Reserves
in Hesse (Germany)
Christoph Muster&Theo Blick
doi: 10.5431/aramit5006
Abstract. In 1990 the Federal State ofHesse (Germany) started a long-term faunistic research programme in se-
lected Strict Forest Reserves. Herewe reportthe results ofthe inventoryofpseudoscorpionsfrom seven reserves:
Goldbachs-undZiebachsrück,Hasenblick,Hohestein,Kinzigaue,NiddahängeeastofRudingshain,Schönbucheand
Stirnberg.Atotalof4567specimens(315-1314persite)belongingto 13species(4-9persite)wererecordedusinga
broad spectrumofmethods.Thebulkofmaterialcomesfrom pitfalltraps(83%),followed byvarioustypesofstem
eclectors(16%).Thestructureandqualityofthespeciesassemblageintheoak-dominatedflood plainforestofthe
Kinzigauediffersstronglyfromthoseoftheremaining reserves,whicharealldominated bybeech.IntheKinzigaue
aspecies-richcommunity(9species)with balanceddominancestructure(Shannon-Index 1.67) hasbeen recorded.
Atthissite,rareormoderatelycommonspeciesreachabundanceproportionsofmorethan50%.Mostimportantly,
twospecieswithstrongaffinitiestopristineforests,DendrochernescyrneusandCherriescimicoides,wereexclusively
recordedfromthisreserve.Incontrast,thebeech-dominated reserveswerestronglydominated byasinglespecies,
Neobisiumcorcinoides (59-91 %ofthe specimens, Shannon-Index < 1), and the abundance proportion ofthevery
commonspeciesexceeded90%ineachoftheseareas.Temperatureandaltitudedeterminethecompositionofthe
speciescommunitiessignificantly.Ofthe recorded species,twowill becategorized intheforthcoming Red Listsof
pseudoscorpionsofGermanyandHesse.Dendrochernescyrneus(cat.3"vulnerable")reachedanabundancepropor-
tion of20% in the reserve Kinzigaue. Dinocheiruspanzeri(cat. G"indeterminate risk") has been recorded in single
specimensfromthereservesHasenblick,Stirnbergand Niddahänge.Thepreferred habitatformostofthespeciesis
theforest.Specieswithstrongaffinitiestoforestsmakeupmorethan95%ofthespecimensinallreservesexceptfor
Kinzigaue,wherethisproportionwas89%.SpeciesrichnessandabundanceswerenotdifferentbetweentheStrict
ForestReserves(managementwasstoppedattheendofthe1980's)andthereferenceareaswithcontinuedforestry.
However,theproportionsofrareand moderatelycommon specieswere higherin theStrict Reservesascompared
tothereferenceareas.
Keywords:Central Europe,Fagussylvatico,natureconservation,oldgrowthforests
Zusammenfassung. Pseudoskorpione (Arachnida: Pseudoscorpiones) aus hessischen Naturwaldreservaten.
In ausgewählten hessischen Naturwaldreservaten (NWR) werden seit 1990 zoologische Langzeituntersuchungen
durchgeführt. Es werden die Ergebnisse zu den Pseudoskorpionen aus sieben hessischen NWR vorgestellt: Gold-
bachs- und Ziebachsrück, Hasenblick, Hohestein, Kinzigaue, Niddahänge östlich Rudingshain, Schönbuche und
Stirnberg.Insgesamtwurden4567Individuen(315-1314proGebiet)aus13Arten(4-9proGebiet)miteinembreiten
Methodenspektrum erfasst. Der Großteil des Materials stammt aus Bodenfallen (83 %) und verschiedenenTypen
vonStammeklektoren (16%).DieStrukturundQualitätderArtengemeinschaften unterscheidetsichsehrstarkzwi-
schendemStieleichen-dominierten HartholzauwaldderKinzigaueundden Buchenwald-Naturwaldreservaten (alle
übrigenGebiete).ImNWRKinzigauewurdeeineartenreicheZönose(9Arten)mitausgeglichenerDominanzstruktur
angetroffen (Shannon-Index 1,67).Seltene und mäßig häufigeArten erreichen hierIndividuenanteilevon mehrals
50%,vorallemaberwurdendieanursprünglicheWäldergebundenenArtenDendrochernescyrneusundChernesci-
micoidesausschließlich hierfestgestellt.ImGegensatzdazuwerdendieBuchenwald-NWRsehrstarkvonNeobisium
corcinoides dominiert (59-91 % der Individuen, Shannon-Index < 1) und der Individuenanteil sehr häufiger Arten
liegtindiesen sechsGebieten bei >90%.Temperaturund HöhenlagederGebietezeigen signifikanten Einflussauf
dieZusammensetzungderArtengemeinschaften.Zweidernachgewiesenen Pseudoskorpion-Artenwerden inden
Roten Listen Deutschlands und Hessens (Arbeitsversion) gelistet sein. Dendrochernes cyrneus (Kat. 3„gefährdet")
erreichtim NWRKinzigaueeinen Individuenanteil von 20%.Dinocheiruspanzeri(Kat.G„Gefährdung unbekannten
Ausmaßes") wurde in Einzelexemplaren in den NWR Hasenblick, Stirnberg und Niddahänge erfasst. Die Mehrzahl
der nachgewiesenen Pseudoskorpion-Arten besitzt ihren Verbreitungsschwerpunkt im Wald.Waldarten im enge-
ren Sinne erreichen in allen Untersuchungsgebieten Individuenanteile von > 95 %, außer im NWR Kinzigaue, wo
derAnteil bei 89% liegt. Die Ende der 1980eraus der Nutzung genommenTotalreservate unterscheiden sich von
weiterhin bewirtschafteten Vergleichsflächen hinsichtlich derArten- und Individuenzahlen kaum. Die Arten- und
Individuenanteile seltener und mäßig häufigerArten sindjedoch in denTotalreservaten deutlich höherals in den
Vergleichsflächen.
5
38 CMuster&TBlick
The knowledge on the distribution and the prefer- diversity-inventory“ approach (Muster 2009, 2013).
red habitats of pseudoscorpions in Germany is lo- Herewepresenttheresults from 7reserves (5 ofthem
wer than in other arachnid groups, i.e. spiders and have an adjacentmanaged site = reference area).With
&
harvestmen (Muster Blick in press). This is re- a total of 4567 specimens this is the second most
flected in the comparatively low numbers of grid comprehensivefaunisticandecologicalstudyonpseu-
squares with records per species,which is on average doscorpions inGermany,outnumberedonlybyDrog-
21 forpseudoscorpions, 105 for harvestmen and 141 la &cLippold (2004,ca.23000 specimens).
for spiders (Staudt 2014, based on 1:25000 maps
with a total of3000 grid squares in Germany).The Materialandmethods
group is still rarelyused in applied studies,biodiver- Studysites. Pseudoscorpions were determined from
sity projects, etc. Nevertheless, the group contains a seven Strict Forest Reserves in Hesse. Data for the
fairly high number ofspecialized species ofcertain sites are listed in Tab. 1, the geographical position
endangered habitats (e.g. bogs, nearly natural fo- within Hesse is shown in Fig. 1.
rests), but in particular ofspecific habitat structures
(bark, rotten wood, nests ofbirds or ants, compost GZ - Goldbachs- and Ziebachsrück(withmanaged
heapsorbarns),resultinginahighpotentialvaluefor reference area)
bioindication in nature conservation.Thiswas consi- HB - Hasenblick (with managed reference area)
deredwhenAnthrenochern.esstellaeLohmander, 1939 HO- Hohestein (with managed reference area)
was included in Annex 2 ofthe Habitats Directive,
where it represents (together with several species of
beetles) the communities ofdecomposition stages in
old growth forests (Drogla2003).
This contributionpresents results offaunistic and
ecologicalresearchin StrictForestReserves in Hesse
(Germany). Currently27 species ofpseudoscorpions
&
are known from Hesse (Muster Blick in prep.).
Systematic surveys in the county were rare (e.g.,
Helversen 1966,Jost 1982) until 1990,when the Fe-
deral State ofHesse started a faunistic inventory in
the Strict Forest Reserves, which is unique and has
no parallelin anyfederal state ofGermany.In Hesse
31 StrictForest Reserves have been established since
1988,with a total area of1228 ha. In the long term
notonlyeverysingle reserve shallbe investigated in-
tensively, but also the succession shall be documen-
ted by repeated inventories (Dorow et al. 1992).The
Senckenberg Research Institute executes this zoolo-
gical research program.The quality ofthis project is
increasedthroughtheinvestigationofmanagedrefe-
rence areas (with continued forestry) in22 sites.
Pseudoscorpions are not included in the standard
set of7 animal groups that are analysed at all levels
&
ofthe project (see Dorowet al.2010,Dorow Blick
2013),butwere addedinthesenseofan„all-taxa-bio-
Fig. 1: Geographical position ofthe investigated Strict Forest
ReservesinHesse/Germany(acronymsinrectangles);theremai-
Dr.ChristophMUSTER,Neukamp29,18581 Putbus,Germany; ningsitesofthe31 StrictForestReservesinHessearemarkedas
E-mail:[email protected] blackdots(afterDorowetal.2010a).Shadedareas:forest
TheoBUCK,Senckenberg,TerrestrischeZoologie,Naturwaldreservate, Abb. 1: Lage der bearbeiteten hessischen Naturwaldreservate
Senckenberganlage25,60325FrankfurtamMain,Germany; (Kürzel in Quadraten);die übrigen der31 in Hessen ausgewie-
E-mail:[email protected] senen Naturwaldreservate sind mitschwarzen Punkten darge-
eingereicht11.2.2015,angenommen10.6.2015,online29.6.201 stellt(nachDorowetal.2010a).SchattierteFläche:Wälder
PseudoscorpionsinForestReserves 39
KI - Kinzigaue see Behre 1989,Blick2011).Alltrunkeclectorswere
NI - Niddahänge E Rudingshain (with managed installed on common beech (Fagussylvatica), except
reference area) for the Kinzigaue, where one eclector was on a li-
SB - Schönbuche (with managed reference area) ving ash tree (Fraxinus excelsior) and the others on
ST - Stirnberg oak Quercus robur). Furthermore tent eclectors on
(
stumps and tenteclectors coveringdeadwood (thick
Fieldwork. In the seven reserves the fauna has branches) were used (details see Tab. 2 and Dorow
been recorded with a broad set of trap types and et al. 1992).
trappingmethods overtwo entireyears (inchwinter) Determination and nomenclature. The deter-
CM
in a standardized way (Dorow et al. 1992, 2010a). mination was conducted by using the keys of
Pitfall traps (generally as groups ofthree = triplets), Beier(1963),Leggöcjones (1988),Devore-Scriban-
window traps (different types, see Tab. 2), colour te (1999), Mahnert (2004) and Christophoryovä et
pans (free,notatthe bark) and trunkeclectors atthe al. (2011b). Nomenclature and affiliation to families
dominating tree species as well on living trees as on follows Muster 8cBlick(in press).
standing and lying trunks (with total contact to the Statistics.The similarityofthe species assembla-
ground and partly without) were used. The applied ges ofthe seven forest sites has been examinedusing
trunk eclectors were open at the bottom, thus the the non-metric multidimensional scaling method
fauna running up the trunks is captured (for details (NMDS), based on Bray-Curtis dissimilarities.This
Tab.1:ShortcharacterizationoftheseveninvestigatedStrictForestReservesinHesse.Forabbreviationsofsitesseetextabove
Tab.1:KurzcharakterisierungdersiebenuntersuchtenhessischenNaturwaldreservate.FlächenkürzelwieobenimText
GZ HB HO KI NI SB ST
Yearofconstitution 1988 1988 1989 1993 1988 1988 1997
Sizeofreserve (ha) 31.3 46.0 26.7 18.1 42.0 27.9 71.1
Sizeofreference area 36.8 40.3 24.4 none 31.7 26.5 none
(ha)
German Gridnumber 5025 4917 4726 5819 5421 5523 5526
(TK25)
°N (WGS84,Centreof 50.9301 51.0586 51.2490 50.1427 50.5239 50.4824 50.4897
thereserve)
°E (ditto) 9.8750 8.6356 10.0468 8.9824 9.2036 9.5397 10.0272
Macrochore East Bergisches East Upper East East East
Hesse Land, Hesse Rhine Hesse Hesse Hesse
Highlands, Sauerland Highlands, Plain and Highlands, Highlands, Highlands,
Vogelsberg Vogelsberg Rhine- Vogelsberg Vogelsberg Vogelsberg
and Rhön and Rhön Mainarea and Rhön and Rhön and Rhön
Mountains Mountains Mountains Mountains Mountains
Altitude (m a.s.l.) 300-365 370-486 455-565 110-115 530-690 370-455 690-910
Meantemperature (°C) 8.0 7.2 8.0 9.6 6.7 7.8 6.3
Meanprecipitation (mm) 748 868 759 712 1175 776 976
Dominatingvegetation beech beech beech common beech beech beech
forests on forestson forestson oak-ash forestson forests on forests on
acidicsoils acidicsoils basic and floodplain basicand acidicsoils basicand
calcareous forest calcareous calcareous
soils soils soils
Referencesfor Dorowet unpubl. Flecht- Blicket Flecht- Dorowet unpubl.
zoologicaldata al.(2009, neretal. al.(2012, neretal. al.(2001,
2010b) (2006), 2014) (1999, 2004a,
&
Dorow 2000) 2004b)
Kopelke
(2007)
40 C.Muster&T.Blick
Tab.2:Overviewofthetrapping methodsandtrapnumbers(in brackets:numbersin reserve/referencesite) .Forabbreviationsof
sitesseetextabove.
Tab.2:ÜbersichtzurFangmethodikunddenFallenanzahlen(inKlammern:AnzahlinTotalreservat/Vergleichsfläche).Flächenkürzel
wieobenimText.
GZ HB HO KI NI SB ST
Periodofinvestigation 1994-1996 1999-2001 1994-1996 1999-2001 1990-1992 1990-1992 2004-2006
Pitfalltraptripletts 16 (8/8) 29 (18/11) 12 (6/6) 12 15 (8/7) 12 (6/6) 23
Additionalsinglepitfalls 8 (3/5) 9 (6/3) 7(3/4) 1 (0/1)
traps
Eclectors onlivingtrees 4(2/2) 4(2/2) 4(2/2) 2 4(2/2) 4(2/2) 2
Eclectors onstanding 4(2/2) 4 (2/2) 4(2/2) 2 4(2/2) 2 (2/0) 2
deadtrees
Eclectors onlyingtrunks 1 (I/O) 3 (1/2) 1 (i/o) 1 4(2/2) 1 (I/O) 1
(withtotalcontacttothe
ground)
Eclectors onfreelying 1 (1/0) 1 (I/O) i (i/o) 1 2 (2/0) 2 (2/0) 1
trunks (onlyinsmaller
partswithcontacttothe
ground)
Tenteclectors onstumps 2 (0/2) 2 (0/2) 1 (0/1) 1 (0/1)
Tenteclectorsfilledwith 2 (1/1) 2 (1/1) 2 (1/1) 1 2 (1/1) 2 (1/1) 1
deadwood
Tenteclectors onthe 2 (1/1) 2(1/1)
floor
Colourpancombinations 2 (1/1) 2 (1/1) 2 (1/1) 1 2 (1/1) 2(1/1) 2
(white,blue,yellow)
Windowtraps (1 m2) 2 (1/1) 1 2 (1/1) 2(1/1) 1
Smallerwindowstraps 2 (1/1) 2 (1/1) 2 (1/1) 2 (1/1)
(„Kreuzfensterfallen“)
Windowtraps atthebark 3 (3/0) 7
ranking ordination method projects the distances al. (2011).The allocation ofindividual species to ca-
into two dimensions, with minimisation ofthe dis- tegorieswasbasedonliteraturedataandourownex-
tortion.Theanalysiswasconductedwiththefunction perience (not restricted to this study).The following
’metaMDS” (2 dimensions, 20 start configurations) categories were distinguished (Tab. 3, compare also
in the package “Vegan” (Oksanen et al. 2011) with Blicket al.2012,2014,BlickScDorow2014):
the program “R, version 2.13.1” (R Development F - largelyrestricted to forest
Core Team 2011). The stress value reports the qua- fc - largelyrestricted to closed forest
lity of the ordination. Values <1 represent the true Mfl - prefers forest edges and clearings
relationships precisely.To interpret the axes and as a - occurs inforestandopenland (with the fo-
test ofthe relevance ofenvironmental variables (see cus notin the forest)
Tab. 7) the Vegan function “envfit”was used. Only mm— occurs in forests aswell as in openland
variables (factors),which correlate significantlywith mo - mayoccur in forests,butprefers open land
the data (1000permutations) are shown asvectors in O - largelyrestricted to openland (or anyother
the ordination diagram. type ofnon-foresthabitat)
Associationwithforesthabitats.Theassociation
ofindividual species with forest habitats was classi- Note that this system reflects exclusively the re-
fied with reference to the system proposed for the lationship with forest, notwith other environmental
German vascular plants and mosses by Schmidt et factors,such as moisture etc.
PseudoscorpionsinForestReserves 41
Results lochernes wideri in trunk eclectors on dead standing
A total of 4567 pseudoscorpions were determined trees and Lamprochernes nodosuswith window traps.
from the seven reserves: 1791 males, 1596 females The remaining species were trapped with up to 10
and 1180 juveniles. They belong to 13 species and different methods. These methods, however, show
%
3 families (Tab. 3): Neobisiidae: 4 species, 94 of quite different efficiencies for the recording of in-
% N
the specimens; Cheliferidae: 1 species,0.05 ofthe dividual species: Neobisium carcinoides and simile
% N
specimens; Chernetidae: 8 species, 6 ofthe speci- weretrappedprincipallywithpitfalltraps, sylvati-
mens. The most numerous species, Neobisium carci- cumandPselaphochernesdubiusmostlyineclectorson
%
noides accounts for 86 ofthe specimens. In single living trees, Chernescimicoides Dendrochernescyrneus
, ,
forest sites 4to 9 species (5,5 ± 1,6) and251 to 1314 and Lamprochernes chyzeri mainly in eclectors on
specimens (652 ± 332) were recorded. dead standing trees, Pselaphochernes scorpioides pre-
Trappingmethods.Thelargemajorityofthepseu- dominantlyin eclectors on lying trunks (onlypartly
doscorpions was caught by pitfall traps (3787 speci- with contact to the ground) andDinocheiruspanzeri
mens,82.9%),followedbythedifferenttypesoftrunk mainly with window traps near the bark. Further-
eclectors (734 specimens, 16.0 %) and the window more, specific life cycle stages may prefer different
traps on the bark (27 specimens, 0.6 %).With other vegetation strata. In Neobisium sylvaticum 53 % of
,
methods only single specimens were trapped, with a the specimens in pitfall traps were adults, while in
maximum of5 specimens permethod (Tab.4). trunkeclectors 79 %belongedto nymph stages.This
A
few species were trapped exclusivelyby a sin- illustrates the importance of using a broad set of
gle method: Neobisium simoniwith pitfall traps,Al- methods to register the species assemblages as rep-
Tab.3:Total numbersofspecimensintheseven reservesin Hesseincludinginformationonassociationwithforestsand raritycat-
egory(RC)accordingtoMuster&Blick(in press):vc=verycommon,c=common,me=moderatelycommon,r=rare.SeeMaterial
andmethodsforabbreviationsofforestassociationandsitenames
Tab. 3:Anzahl erfasster Individuen in sieben hessischen Naturwaldreservaten und Einstufung hinsichtlichWaldbindung und Be-
standskategorien(RC)nachMuster&Blick(inpress):vc=sehrhäufig,c=häufig,mc=mäßighäufig,r=selten.ErklärungderAbkür-
zungenfürWaldbindungundFlächenkürzelinMaterialundMethoden
Taxon GZ HB HO KI NI SB ST 2 Association RC
withforests
Neobisiidae
Neobisiumcarcinoides(Hermann,1804) 750 1134 547 46 439 278 372 3566 F vc
Neobisiumsimile(L.Koch,1873) 22 149 114 15 300 F VC
Neobisiumsimoni(L.Koch,1873) 40 40 F c
Neobisiumsylvaticum(C.L.Koch,1835) 12 50 32 187 5 98 384 F VC
Cheliferidae
MesocheliferressliMahnert,1981 1 1 2 F c
Chernetidae
Allocherneswideri(C.L.Koch,1843) 1 1 F c
Chernescimicoides(Fabricius,1793) 87 87 F mc
Dendrochernescyrneus(L.Koch,1873) 49 49 F r
mm
Dinocheiruspanzeri(C.L.Koch, 1837) 8 1 4 13 c
Lamprocherneschyzeri(Tömösväry,1882) 1 3 4 fl r
Lamprochernesnodosus(Schrank,1803) 1 1 2 mo c
Pselaphochernesdubius(O.P.-Cambridge,1892) 37 2 5 16 60 fl mc
Pselaphochernesscorpioides(Hermann, 1804) 18 10 27 1 3 59 mo c
Specimens 827 1314 600 251 743 315 517 4567
Species 4 6 4 9 6 5 5 13
Shannon-diversity 0.41 0.49 0.32 1.67 0.97 0.49 0.81
1
I 1 11 I 1
42 C.Muster&TBlick
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PseudoscorpionsinForestReserves 43
resentatively and completelyas possible (see Dorow higher in the unmanaged areas (reserves),whereas
et al.2010a). in Hasenblick and Schönbuche it was higher in
Comparison of the reserves with their man- the managed reference areas, while no differences
aged reference areas. The quality of this research were observed in Goldbachs- und Ziebachsrück
A
programis increased through the inclusion of22 ad- (Fig. 2). higher number of individuals were
jacent managed sites/reference areas, where forestry trapped in the reserves ofHasenblick and Hohe-
is continued. Five ofthe seven reserves included in stein compared to their reference areas, and lower
this study have such reference areas (not present in numbers inGoldbachs- und Ziebachsrück,Schön-
Kinzigaue and Stirnberg). buche and Niddahänge (Fig. 3). Note that the re-
Neither the species numbers nor the numbers cordingschemewas notcompletelyidenticalinthe
of pseudoscorpion individuals were significantly reserves and the correspondingreference areas (see
different between the reserves and the reference Tab.2).The number ofpitfall trap sites reflects the
areas (Mann-Whitney U-Tests, p > 0.05). In Ho- number ofdifferent structures in the reserves and
hestein and Niddahänge the species number was the reference areas.
Fig.2:Comparisonofthe
species numbers of the
pseudoscorpions in the
7 Strict Forest Reserves
(SFR) and the 5 reference
sites(REF)inHesse
Abb. 2: Vergleich der
Artenzahlen von Pseu-
doskorpionen in Total-
reservaten (SFR) und
deren Vergleichsflächen
(REF) von 7 hessischen
Naturwaldreservaten GZ HB HO Kl NI SB ST
800
Fig.3:Comparisonofthe
specimennumbersofthe
pseudoscorpions in the
7 Strict Forest Reserves
(SFR) and the 5 reference
sites(REF)inHesse
Abb.3:Vergleich derAn-
zahl erfasster Individuen
von Pseudoskorpionen in
Totalreservaten(SFR)und
deren Vergleichsflächen
(REF) von 7 hessischen
Naturwaldreservaten GZ HB HO Kl NI SB ST
44 C.Muster&T.Blick
Tab.5:Abundance(standardizedtoindividuals/pitfalltrap)ofpseudoscorpion speciesfrom pitfalltraps in strictforestreservesas
comparedtomanagedreferencesites(KlandSTomittedduetolackofreferencesites).Forabbreviationsofsitesseetextabove
Tab.5:VergleichderstandardisiertenAbundanzvonPseudoskorpionen(Individuen/Bodenfalle)ausBodenfalleninTotalreservaten
(SFR)undderenVergleichsflächen(REF)in5hessischenNaturwaldreservaten(inKlundSTwurdenkeineVergleichsflächenbeprobt).
FlächenkürzelwieobenimText
Strictforestreserves (SFR) ]Referencesites (REF]) all all %
SFR REF SFR
GZ HB HO NI SB GZ HB HO NI SB
Dinocheiruspanzeri 0,09 0,09 0,04 0,033 0,031 51,4
Lamprocherneschyzeri 0,03 0 0,008 0,0
Lamprochernesnodosus 0,04 0,05 0,007 0,008 45,8
Mesocheliferressli 0,04 0,007 0 100,0
Neobisisumcarcinoides 10,7 10,3 14 6,81 6,56 15,9 17,6 10 10,2 8,42 9,887 13,11 43,0
Neobisiumsimile 0,37 2,13 2,22 0,78 0,41 1,03 2,16 0,05 1,327 0,795 62,5
Neobisiumsylvaticum 0,11 1,92 2,74 0,17 0,18 0,19 4,52 0,11 0,86 0,984 46,6
Pselaphochernesdubius 0,74 0,08 0,33 0,59 0,53 0,187 0,213 46,8
Pselaphochernesscorpioides 0,59 0,02 0,04 0,07 0,27 0,12 0,087 58,1
At the species level, however, some differences (German category„G“). In each ofthe reserves Ha-
are recognisable. Neobisium carcinoides was trapped senblick, Kinzigaue, Niddahänge and Stirnberg one
more numerouslyin almostallreference areas,which ofthese two species has been recorded. The reserve
indicates a preference for more open forest stands, Kinzigaue has the highest proportionate abundance
whereas for N simile the reverse pattern emerged (24.3 %) ofRed Data List species (< 1 % in all other
(Tab.5). reserves).
Qualitative characteristics of the assembla- Figs. 4 and 5 show the proportions of species
ges. The assemblages of four beech forest reserves and individuals of rarity categories, as included in
&
(Goldbachs- und Ziebachsrück, Hasenblick, Hohe- the latest Red Data List (Muster Blick in press).
stein and Schönbuche) show a similar structure, as The species were classified into seven criteria-classes
they are strongly dominated by Neobisium carcinoi- (from extremelyrare to very abundant) based on re-
des (> 85 %) and contain a relatively low number of cordsonthe 1:25000gridsquaresofGermany(mean
species (max. 6).This is furthermore reflected in low value about 129 km2 per grid) (Staudt 2014). Extre-
values ofthe Shannon-diversity (< 0.5, see Tab. 3). mely rare and very rare species have not been found
The two beech reserves in the higher low mountain inthereserves.Theverycommon speciesreachabout
range (Niddahänge, Stirnberg) have a comparable 50%,withtheexceptionofKinzigaue,wheretheper-
number ofspecies. But the structure ofthe commu- centage was rather lower (Fig. 4).This is even more
nities is more balanced, with other Neobisium spe- obviousregardingthe abundanceproportion (Fig.5):
cies reaching high percentages, resulting in higher whereas the percentage ofthe individuals ofthevery
Shannon-values (0.8—1.0). Totally different is the common species was higher than 90 % in all other
situation in thelowland floodplain forest,dominated reserves,itwas about 1/3 in the Kinzigaue. Here,the
by common oak (Kinzigaue). Here, the structure of relative abundance ofthe moderately common and
dominance is balanced, the species number is consi- rare species reaches more than 50 %.
derablyhigherand ahighervalue ofShannon-diver- The analysis ofthe pseudoscorpion assemblages
sityis reached (1.67). with respect to their relationship to forest habitats
In the Red D&ataListofthe Germanpseudoscor- results in the (unsurprising) insight, that all recoWr-
pions (Muster Blick in press) two species from ded species are typical forest dwellers (categories
M
the Strict Forest Reserves in Hesse are listed. Den- and including their subcategories) (Tab. 3).More
drochernescyrneusis classified as vulnerable (German than 98 % ofthe specimens belong to species which
category „3“), while the degree ofendangerment of are stronglyassociatedwith foresthabitats (Tab.6: F
Dinocheiruspanzeri could not be classified in detail and fl). Again there is one exception, the Kinzigaue,
PseudoscorpionsinForestReserves 45
%
where 10.8 ofthe specimens belong to Pselapho- ve Kinzigaue has a separatedposition.Axis 2 divides
chernes scorpioides a species that can be found regu- the beech forests on acidic soils (GZ,HB, SB) from
,
larlyinforests,but has its focus outside offorests. those on basic and calcareous soils (HO, NI, ST),
Relationships ofthe species communities. The which are,atthe same time,the threewith the high-
relationships ofthe species communities ofthe se- est elevation. Of the tested abiotic factors, altitude
ven reserves (inch their reference areas) and the in- and the average temperature correlate significantly
fluence ofabiotic variables was examined by means with the position of the reserves in the ordination
ofnon-metrical multidimensional scaling (NMDS). (Tab. 7, Fig. 6). In contrast, an influence ofprecipi-
The NMDS-ordination (stress < 0.1) shows the area tation orgeographicalposition is not detectable.The
Hasenblick in the centre of the diagram (Fig. 6), species Allochernes widen, Cherries cimicoides, Dend-
in nearly equal distance to the other beech forests. rochernes cyrneus and Lamprochernes chyzeri are asso-
NMDS
Alongaxis 1 ofthe theoakdominatedreser- ciated with the lower sites with higher temperature,
Fig. 4: Percentage of
the species of the pseu-
doscorpions of seven
Hessian Strict Forest Re-
serves (reference areas
vc
included), categorised
after their frequency in c
Germany, vc = verycom- COmc
mon, c = common, me =
moderately common, r = r
rare
Abb. 4: Artenanteile der
Häufigkeitsklassen von
Pseudoskorpionen in
sieben hessischen Natur-
waldreservaten.vc=sehr
häufig, c = häufig, mc
mäßighäufig,r=selten
Fig. 5: Percentage ofthe
individuals of the pseu-
doscorpions of seven
Hessian Strict Forest Re-
serves (reference areas vc
included),categorisedaf- c
tertheirfrequencyinGer- mc
many.vc-verycommon,
c - common, mc - mod- EBr
eratelycommon,r-rare
Abb.5:Individuenanteile
der Häufigkeitsklassen
von Pseudoskorpionen in
sieben hessischen Natur-
waldreservaten.vc-sehr
häufig, c - häufig, mc -
mäßighäufig,r-selten
46 C.Muster&TBlick
Tab.6:Associationofthespeciesassemblageswithforestsinthesevenreserves(seetextandTab.3)
Tab.6:WaldbindungderArtengemeinschaftenindeneinzelnenNaturwaldreservaten(vgl.TextundTab.3)
Relationship toforests GZ HB HO KI NI SB ST
F-largelyrestrictedtoforest 93.3 % 98.6% 99.7% 86.1 % 99.6% 94.6% 98.6%
fl-prefersforestedges andclearings 4.5 % 0.1 % 0.3 % 3.2% 5.1 %
mm - occursinforests andopen 0.6% 0.1 % 0.8 %
land
mo - mayoccurinforests 2.2% 0.8% 10.8 % 0.3 % 0.3 % 0.6%
whileNeobisiumsimonishows a relationship with si- al.2008).SeveralspeciesofChthoniidaeareregularly
tes ofhigher elevation. As expected, the ubiquitous trappedwith pitfalls, and the most common species,
Neobisium carcinoides is positioned in the centre of Chthonius (Ephippiochthonius) tetrachelatus, has been
the ordination (Fig. 6). collected with this method in the course ofbiospel-
eologicalsurveysinHesse (Zaenker2001,Reissetal.
Discussion 2009). In the forest ofEttlingen (Baden-Württem-
This study ofseven Strict Forest Reserves in Hesse berg) C. tetrachelatusmade up 24% ofthe specimens
(including five reference areas) resulted in records caughtwithpitfalls (Braun &cBeck 1986) and in the
of 13 species ofpseudoscorpions. This is equivalent Luxembourgian forest“Schnellert”even four species
to 26 % of the 50 species known from Germany ofChthoniushavebeen found (Groh2007).
(Muster<5cBlickinpress) and50%ofthe26 species Due to the low overall number of species we
&
known from Hesse (Muster Blickin prep.).These refrained from using rarefaction methods to assess
ratios are higher than the average in other groups of the completeness ofthe inventory. But the intensity
animals (23 % resp. 32 %,Dorowet al.2010a).There ofthe sampling and the broad spectrum ofapplied
are great differences in the representation of the methods implya rather comprehensive survey.There
more species-rich families among the pseudoscorpi- exist only few regional faunistic studies on Central
ons. Whilst 33 % of the German and 80 % of the European pseudoscorpions of similar magnitude.
Hessian Neobisiidae and 53 % ofthe German and Comparedto othersites,the nine species inthe Kin-
%
73 ofthe Hessian Chernetidae have been found, zigaue is a fairly high number, and even four to six
the species-rich family Chthoniidae (14 species in species in the other reserves are not unusually low.
Germany,5 species inHesse)waslackingcompletely Some examples are given to illustrate this: Strebel
in the collections from the Strict Forest Reserves in (1961) collected four species in the “Siebengebirge”
&
Hesse.With regard to the high intensityoftrapping (North Rhine-Westphalia), Helversen Martens
and the broad spectrum ofsurvey methods this lack (1971) listed nine species from the gorge of the
ofChthoniidae remains unexplainable. In fact, it is “Wutach” (Baden-Württemberg), Drogla (1977) re-
known that some species of Chthoniidae show ag- cordedfive species in amixeddeciduous forestinthe
gregated distributions (sometimes with high local low mountain range ofthe nature reserve “Tiefental
&
abundances, see Drogla Lippold 2004, Muster et bei Königsbrück”(Saxony),an intensive inventoryin
nearly natural forests in Luxembourg revealed the
Tab.7:Correlationofthevariables/factorswiththeNMDSordi- presence of five or six species (Groh 2007, Köhler
nation,significantfactorsareinbold et al. 2011), the collections byJost (1982) in the na-
Tab.7:KorrelationvonEinflussvariablen mitderNMDSOrdina- ture park“HoherVogelsberg”(Hesse) yielded eleven
tion,signifikanteUmweltvariablensindfetthervorgehoben species; the same number as found by Stählavsky
&
Variable/Factor Axis1 Axis2 r2 P(>r) Kräsny (2007) in the valley of the lower Vltava
Altitude(m) 0.66598 -0.74597 0.8936 0.008** (Czech Rep.). On the other hand, Drogla (1988)
Temperature -0.88388 0.46772 0.8638 0.013* found in a forest near “Deutsch Paulsdorf” (Upper
Precipitation 0.85473 -0.51907 0.6253 0.139 Lusatia,Saxony) onlyasingle species (Neobisiumcar-
cinoides) and 587 specimens from a floodplain for-
“East 0.20875 -0.97797 0.1286 0.758
est at the river Rhine (Rhineland-Palatinate) (Marx
“North 0.47475 -0.88012 0.0572 0.875 et al. 2008) as well as 3777 specimens in the forest