Vertical distribution and abundance of pseudoscorpions (Arachnida) in the soil of two different neotropical primary forests during the dry and rainy seasons

10 Pages·1993·1.9 MB·

by  Joachim Adis

VERTICALDISTRIBUTION ANDABUNDANCEOFPSEUDOSCORPIONS
(ARACHNIDA) INTHE SOILOFTWODIFFERENTNEOTROPICAL PRIMARY
FORESTS DURING THE DRY AND RAINY SEASONS
JOACHIM ADISand VOLKER MAHNERT
Adis,J.andMahnert,V. 1993 11 11:Verticaldistributionandabundanceofpseudoscorpions
(Arachnida) in thesoiloftwodifferentneotropicalprimary forestsduringthedry andrainy
seasons.MemoirsoftheQueenslandMuseum33(2): 431-440. Brisbane. ISSN0079-8835.
Pseudoscorpionswereextractedfrom0-14cmsoildepthintwodryland(upland)forestsnear
Manaus, Brazil. In a primary forest on yellow latosol, about 1700 specimens per m were
obtainedduringthedryseasonand 1135duringtherainyseason.Theyaccountedfor 10-15%
ofall arthropods extracted, excluding Acari and Collembola. In a primary forest on white
sand soil (campinarana), about530 specimens perm" were obtained duringthe dry season
and480during the rainy season.They accountedforonly3-4% ofall arthropodsextracted,
againexcluding Acari andCollembola. Significantbutdifferentcorrelations were found in
both forest types between the abundance of pseudoscorpions and changing moisture,
temperature and pH conditions in relation to soil depth and season. Neither during thedry
seasonnorduringtherainyseasonwastheabundanceofpseudoscoq^ionsinmineralsubsoils
higherinresponsetothechangingsoilmoisturecontentinorganic layers.Thiswasreported
forarthropodsfromforestsintheseasonaltropicswhereperiodswithoutprecipitationoccur,
Results are discussed at the species level. They are compared with published data on the
vertical distribution and abundance ofpseudoscorpion species from the yellow latosol ofa
secondary dryland forest (dry season and rainy season) from the sameregion.
In zwei Festlandwaldern in der Umgebung von Manaus wurden Pseudoskorpionc aus
0-14cm Bodentiefe extrahiert. In einem Primarwald auf gelbem Latosolboden wurden
wahrend derTrockenzeit 1700 und wahrend derRegenzeit 135 Individuen pro m" nach-
1
gewiesen.Sicrcprasentierten 10-15%allerextrahiertenArthropoden,AcariundCollembola
ausgenommen. In einem Primarwald aufWeiBsandbodcn (campinarana) wurden wahrend
derTrockenzeit 530und wahrend derRegenzeit480Individuen pro m nachgewiesen. Sie
reprasentierlennur3-4%allerextrahierlen Arthropoden,Acari undCollembolen wiederum
ausgenommen. Signifikante aber unlerschiedliche Korrelationen crgaben sich in beiden
Waldtypen zwischen der Abundanzder Pseudoskorpionc unddersich andernden Feuchle.
derTemperatur und dem pH in Bezug aufBodentiefe und Jahreszeit. Weder wahrend der
Trockenzeit, noch wahrend der Regenzeit, war die Abundanz der Pseudoskorpionc im
mineralischen Unterboden als Folge aufdie sich andernde Fcuchte im organischen Ober-
boden hoher. Dies wurde furArthropoden in Waldern dersaisonalenTropen, dieTrocken-
perioden durchlaufen, nachgewiesen. Die Ergebnisse werden aufArtniveau diskutiert. Sie
werdenmit publizierten Daten iibcrdie Vcrtikalvcrteilung und Abundanzvon Pseudoskor-
pionartenauseinemSekundarwaldaufgelbemLatosolbodenimgleichenGebietverglichen.
QPseudoscorpiones, abundance, seasonality, verticaldistribution, Neotropics.
Joachim Adis, Tropical Ecology Working Group, Max-Planck-Institute for Limnology,
Postfach165,D-2320Ploen, Germany,incooperationwithNationalInstituteforAmazonian
Research(INPA),Manaus,AM,Brazil; VolkerMahnert,Museumd'Histoirenaturelle, Case
Postale434, CH-1211 Geneve6, Switzerland; 23November1992.
In those wet but markedly seasonal tropics upland) forests experience a rainy season
where periods without precipitation occur, ter- (December-May; average monthly rainfall 211-
restrial arthropods are reported to migrate to 300mm) and a 'dry' (= drier) season (June
mineral subsoils in the dry season as a response November; average monthly rainfall 42-
to changing humidity in organic layers (Beck, 162mm). Annualprecipitation is2105mm(based
1964; Bullock, 1967; Goffmet, 1976; Lawrence, on 75 years of records from the meteorological
1953; Levings and Windsor, 1984; Liebermann station at Manaus, cf. Ribeiro and Adis, 1984).
and Dock, 1982; Merino and Serafino, 1978; About 75% ofthe rainfall (1500mm) is recorded
Petersen and Luxton, 1982; Rybalov, 1990; during the rainy season. This had no observable
Strickland, 1947; Willis, 1976 and others), difference in vertical distribution of terrestrial
Centrai Amazonian dryland (= non-flooded arthropods in primary forests on yellow latosol
.
432 MEMOIRS OFTHEQUEENSLANDMUSEUM
DRY SEASON RAINY SEASON
N/m 702.5 635.1 259.8 105.9
45 -|(%] r/oj
2
N/rr/=1,703 3 2: N/m -1,135.7
30 30 -
•-
15 - 15 **r
r
0-3-5 3.5-7.0 70-10.5 10.5-UO 0-35 35-7.0 70-10.5 10.5-U.O
Soil Depth (cm) Soil Depth (cm)
—
|=nymphs Uodults • • = soil moisture content
I
FIG. 1.Distributionofpseudoscorpionsin soil andsoilmoisturecontent(bothin%).Samplestakenevery3.5cm
todepth of 14cm in dry season and in rainy season in primary dryland forestonyellow latosol nearManaus,
Brazil. Total catch each season = 100%. Abundance for each soil layer (N/m ) and for total catch/ season
(SN/m2
).
and on white sand soil (Adis et al, 1989a, b, thaler(1956),Prance(1990),Prancedal. (1976)
unpublished data; Morais, 1985). However, and Rodrigues (1967) provide detailed botanical
results were based mainly on data for orders. descriptions oftheforest. Microclimatic dataare
Evaluation of sampling data at the species level given by Decico et al. (1977), Marques et al.
is now possible forpseudoscorpions, since iden- (1981) and Ribeiro and Nova (1979). The soils
tification has been completed by the second neartheDuckeForestReserveweredescribedby
author. Falesi and Silva (1969) as deep, yellow latosols,
strongly weathered, excessively to very strongly
STUDY AREAS acidic, of heavy texture in all profiles (A-C: 0-
1 10cm) and with clay content of the B horizon
PrimaryForeston Yellow Latosol varying from 50-70%. In the study area, where
entomologicallong-terminvestigationshavepre-
Sampling was carried out during the rainy
viously been undertaken (Adis and Schubart,
(March) and dry seasons (October) of 1987 in a
dryland terra firme forest at the Ducke Forest 1984; Morais, 1985; Penny and Arias, 1982), the
Reserve (= Reserva Florestal A. Ducke, 2°55'S, soil carried a l-3cm thick humus layer (A ), in-
59°59'W) of the National Institute for terspersed with fine roots and a thin leaf litter,
Amazonian Research (INPA, Manaus), situated covering most ofthe surface. For further details
oPnentnheyMaanndauAsr-iIatsa,co1a9t8i2a)r.aThhieghawraeay(saAmMp-l0ed10;wacfs onDtuhreinsgtutdhyear1e9a87sereaiPneynnsyeaasonnd,A3ri2a2s,mm198o2f.rain
classified as high terra firme forest (Takeuchi, were recorded in March (data from the
1961; cf. Brinkmann, 1971) with several species meteorological station at Ducke Forest Reserve,
of large, broad-trunked trees (with or without provided by M. de N.G. Ribeiro at INPA,
buttresses). The trees reached 44m in height Manaus). On the day of sampling (March 20,
(average height: 22m) and formed a closed 1987) soil moisturecontent (= weightdifference
canopy.Approximately235species,representing between wet and dried soil samples in%) was
43 families of trees, were recorded in the study 20.0% at 0-3.5cm (= humus layer), 17.5% at
area. Most frequent were Leguminosae, 3.5-7cm, 15.5% at 7-10.5cm and 17.7% at 10.5-
Rosaceae, Lauraceae, Sapotaceae and 14cm soildepth, respectively (=mineral subsoil)
Lecitidaceae.Theforesthadapatchyunderstorey (Fig. 1). Soil temperature at 10 a.m. decreased
andascantyherblayer.Guillaumet(1987),Lech- from 25.2°C in the top 3.5cm to 24.5°C at a soil
1
NEOTROPICALPSEUDOSCORPIONS
DRY SEASON % N/rrT RAINY SEASON % N/rrV
IOOjD tOUO 1,135.7
—esk::-:.-:-»;-:-=M 1ndens,_-_-;;'--. ^6a.j:--i 1,159.7 3516.E
nra
7 minor 12 - . i 1 T *T111C*
•••.•••J e.a 6 t.roivpl B1.B
1 schusie.i A grocfl ,: IB 33.7
S3 4 grccilis 3.7 62.4 i sctju^ton 19.2
] P. homodentotus o.e |*.J I tenuis
I 1 6i0 2I0 4i0 50 aoit%]
FIG.2.Dominancet'ft.landabundance(N/m2)ofspeciesofpseudosaifpiimsextracted Irara Im:ofsoil(0-
dtaetpotsho)ldnuerairnMgadnrayusse,aBsroanz(iOlcTtootbaelrc1a9tc8h7)oaf-nedacrhaisneyasSoenas=o=n1(0M0a%r.c(hSe1e98t7e)xtifnorpfruirmtahreyredxrpyllaannadtifoonr*e.st on yf
depthof 14cm The soil pH increased from 3.3 in pling (March IS. 1988] soil moisture content =
|
the top 3.5cm to 3.8 ill the lower layers (3.5.- weight difference between wet and dried sod
14cm). During the IM87 dry season. 139mm of samples in %) increased from 20.1% in the top
rain were recorded in October (data from the 3.5cmto26.2% at 10.5cmdepth (= humus layer'
meteorological station atDuckeForest Reserve) and dropped to 9 9% at 10.5-I4cm (- white sand
On the day of sampling (October 14, 1987) soil soil) (Fig. 4). Soil temperature at 1) a.m
moisturecontentincreasedfrom 13.6% inthetop decreasedfrom 24.7°Cinthetop3.5cmto24.5'JC
3.5cm to 19.1% at a soil depth of 14cm (Fig )) at a soil depth of 14cm. The pH ofthe soil was
and the soil pH increased from 3.3 lo 3.8. Soil 3.2 (0-3.5cm), 3.3 (3.5-7cm), 3.6 (7-!0.5cmiand
t24e.m7pDeCr)atfurroemattheJs1uraf.amc.e tvoar1i4ecdm.slightly (24.6- d3.r7y (s1e0a.so5n-,147cm7im,mreospfecratiinvfeallyl. wDeurreinrgectohrede1d in
August (data from the meteorological station of
EMBRAPA/UEPAH, km 54). Onthedayofsam-
PrimaryForest on White Sand Soil
pling (August 05. I9S8] soil moisture content
Sampling was carried out during the rainy decreased from 21.9% in the top 3.5cm to $
season (March) and the dry season (August) of at 10.5cm depth (= humus layer] and to 9.3% al
1988 in a dryland campinarana forest of the 10.5-14em (= white sand soil, Fig 4). Soil
Biological Reserve INPA/SUFRAMA (approx. temperature at 3 p.m. decreased from 27.1°C in
2°30'S,60°I0,W). atkm45 (formerly km 62) on the top 3.5cm to 2S.5°C at a soil depth of 14cm.
the ManausBoa Vista highway(BR-174). Cam- The pH of the soil was 3.6 (0-3.5cm), 3.5 (35-
pinarana (= caatinga arborea) was classified as a 7cm). 3.5 (7-10.5cm) and 3,6 (10.5-I4cm>.
elati\ely lightforestonwhitesandsoil with respectively. The relative light intensity on the
thin-stemmedtrees 10-20mhigh,withoccasional forest floor was about 2 \% (210 lx; comparative
large, broad-trunked individuals, with orwithout valueintheopenairat3 p.m., 10.000lx (overcast
buttresses,a patchyunderstorev andnoherblayer sky);cf Brinkmann, 1970, 1971).
(Guillaumet, I9S7, Anderson, 1981; Lisb>>a.
1975). Data on geomorphologv and soil genesis METHODOLOGY
are found in Chauvcl etal. ( 1987) Floral inven-
tories have been given by Lisboa (1975). Braga
(1979), Anderson etal i 1975). Anderson ( i9sT) In bothstudy areas, sixsoil samples were taken
and Guillaumet (1987), while the microclimatic along a transect al random intervals with a split
datawereprovidedby RibeiroandSantos ( 1975). corcr (a steel cylinder with lateral hittj
In the study area, the white sand soil carried a diameter 2 Iem, length 33cHl) which was driven
humus layer that was 10-1 tern thick (Ao), intothesoilwith amallet.Each samplewastaken
penetrated by a matting of roots and a thin? sur- toadepthof14cm and wasthen divided intofour
face-covering leaf litter. subsamples of 3.5cm each. Animals were ex-
During the 1988 rainy season, 293mm of rain tracted from subsamples following a moditk-
were recorded in February and 2K0mm in M- method of Kempson (Adis, 1987). All pseudn
(data from the meteorological station of scorpions were separated by species [adults
EMBRAPA/UKPAK, km54).Ontheday ofsam- (males and females), nymphai insiars| and iheil
r r
434 MEMOIRS OFTHEQUEENSLANDMUSEUM
DRY SEASON RAINY SEASON DRY SEASON RAINY SEASON
N/m : 105.9 144 4.8 N/m : 168.438.5 4JB 1107 28.9
100 Id eobisium schusteri 100 -i Tyronnochthonius minor
•2. N/rrf= 192 ZN/m2=211.7 i—i ^N/m2=139.6
-
50 50
E —
\—\
i i r
N/m': 1347 H4 77.0 4.8 N/m2:231.0 5678255j01055 2262264.7235.8129.9
100 -, Brazilatemnus browni 100 Microblothrus tridens
2 N/m =81.8 3: N/rrT= 1.159.7 ^ N/rrr= 856.6
50 -
BH
fl
r~n t r i r
-3.5-7.0 -1Q5-KJ0 -3.5 -7.0-10.5-14.0
N/m : 48.1 14.4 4.8 24.1 4.8 Soil Depth (cm)
100 -,
Albiorix gracilis
— X N/m*= 625 2: N/m = 33.7 f | = nymphs ] = adults
. i
50 -
lii—n—
I | 1 1 r^ r i i i
3.5 -7.0-10.5 -14.0 3.5-7.0-10.5-14.0
Soi Depth cm
I ( )
FIG.3. Verticaldistributionoffivespeciesofpseudoscorpions in soil (%). Samplestakenevery 3.5cm todepth
of 14cmduringdry season andrainy season in primary dryland forest onyellow latosol nearManaus, Brazil.
Totalcatcheach season = 100%. Abundanceforeachlayer(N/m2)and totalcatchperseason (S N/m2).
abundance was calculated for 1 m2. Vertical dis- disregarded) in the study area (Adis et al, un-
tribution of adults and nymphs in relation to published data). During the dry season, about
changing conditions of soil moisture content, 41% of all 1,700 (±71) pseudoscorpions were
temperature and pH was statistically evaluated collected from the top 3.5.cm, 37% from below
with the linear correlation test (Cavalli-Sforza, the humus layer (3.5-7cm) and 22% at 7-14cm
1972), usingthe original field data. depth. About 54% of all specimens collected
represented juvenile stages (Fig. 1: nymphs).
RESULTS Decreasing abundance of pseudoscorpions at
greater soil depths was significantly correlated
PrimaryForeston Yellow Latosol with increasing soil moisture content (adults and
Pseudoscorpions accounted for 10-15% of all nymphs: P<0.01, r = -0.998, nymphs only:
arthropodsextractedfrom 1m soilof14cmdepth P<0.05, r = -0.956; n = 4). During the rainy
(10,000-12,000 ind./m2, Acari and Collembola season, vertical distribution was similar(Fig. 1):
—
. )
NEOTROPICALPSEUDOSCORPIONS 435
DRY SEASON RAINY SEASON
255D U43 62A 7Z1 N/m2; 351.2 53.C) 33.7 A3.3
60-
2
2 N/rn = 533.8 2N/m =461.2
45J
30-
-•^"
-
15
•
~
0-35 35-7.0 7.0-10.5 10.5-UJO 0-35 3.5-70 7.0-10.5 105-K.0
Soil Depth (cm) Soil Depth (cm)
= nymphs I l = adults = soil moisture content
FIG.4.Distributionofpseudoscorpionsinsoilandsoilmoisturecontent(bothin%). Samplestakenevery3.5cm
to depthof 14cmduringdryseasonandrainy seasoninprimarydrylandforestonwhitesandsoil nearManaus.
Brazil.Totalcatcheachseason= 100%.Abundanceforeachsoillayer(N/nr)andtotal catch/season(SN/nr
38% of the 1.100 ( ±46) pseudoscorpions were catch.Lessabundant(>10%)wereMiratemnidac
recovered from the top 3.5 cm, 29% from 3.5- (Brazilaiemnus browniMuchmore ; dry season:
7cm and 33% from 7-14cm soil depth. About 149±20 ind./m2, rainy season: 82± 13 ind./m2)
61% ofthe total catch were nymphs. Decreasing and Ideoroncidae (Albiortx gracilis Mahnert).
abundance of pseudoscorpions at greater soil Few differences in dominance were found for
depthswassignificantlycorrelated with decreas- species captured during both seasons, however
ingsoiltemperature(adultsand nymphs: P<0.01 theirabundance varied (Fig. 2).
r= +0.993. nymphs only: P<0.05, r=+0.956; n= Except for A. gracilis, vertical distribution of
4). pseudoscorpions in the soil differed between
Six species ofpseudoscorpions were collected species but not within species when comparing
in both seasons (Fig. 2). All were previously different seasons (Fig. 3). /. schusteri and B.
reportedfromsoilsofAmazoniandrylandforests browniwereonlyfoundat 0-7cmsoildepth, with
(Mahnertand Adis, 1985). Syarinidae were most abundancesbeing greatest in the top3.5cm. This
abundant, with Microblothrus tridens Mahnert is also true for 7. minor which lived, like A.
(dry season: 160±55 ind./m*, rainy season: gracilis, to a soil depth of 10.5cm. However,
1
860±24 ind./ra2), IdcobisiumschusteriMahnert occurrence was restricted to the upperA7cm in T.
and Ideoblothrus tenuis Mahnert accounting for minorduring the rainy season and in gracilis
74-78% of the total catch (Fig. 2>. Chthoniidae duringthedryseason (Fig. 3). /. tenuiswasfound
was the next most abundant family, with Tyran- only at a soil depth of 3.5-7cm (= below the
nochthonius (T.) minor Mahnert (= Lagynock- humus layer) during both seasons, whereas M.
thoniusminor(Mahnert)):dryseason: 212± 17.4 tridens occurred in all soil layers (0-l4cm).
ind./m2, rainy season: 140± 14ind./m2) and (dry Duringthedryseason,significantcorrelationwas
season only) Pseudochthonius homodentatus observed between decreasing abundance of T.
Chamberlin representing 12-13% of the total minor and greater soil depths (Fig. 3) and the
1 Atthis locality, B. browniisapparently smaller. It mayhaveadaptedto itsenvironment, thus representing an
c-co-species.
436 MEMOIRS OFTHEQUEENSLANDMUSEUM
DRY SEASON % N/m RAINY SEASON % N/m
100.0 533.8 100.0 481.2
|^3.3 230.9 35.0 158,4
i4-.-.--.. 30_6 163.6 32.0 154.4
B.browni 17. ] 91.3 -:-:=:-;-z:-;fi;ijM. tridens 28.0 134.8
I, schusten 5.4 28.8 I schusten 3.0 H.4
R homodentatus 1.8 9.5 A. arboncola 1.0 4.8
A. arboncola 0.9 4.8 T rolundimanus 1.0 4.8
P crassifemoratus 00..99 4.8
"1
20 60 (%) 20 60 (%
FIG. 5. Dominance(%) andabundance(N/m2)ofspeciesofpseudo scorpionsextractedfrom lm2 soil (0-14cm
depth)duringdry (August 1988) andrainy (March 1988) seasons inprimary drylandforeston whitesand soil
nearManaus, Brazil. Total catchofeach season = 100%. (Seetextforfurtherexplanation).
increased soil moisture content (P<0.05; adults all specimens collected represented juvenile
and nymphs: r = -0.959, adults only: r = -0.968; stages (Fig. 4: nymphs). Decreasing abundance
P<0.10;r=-0.936fornymphsonly;n=4,respec- of pseudoscorpions at greater soil depths was
tively). Similar results were recorded from A. significantly correlated with decreasing soil
gracilis (P<0.05; adults and nymphs: r= -0.969, moisture content (P<0.05; adults and nymphs: r
nymphsonly: r=+0.974;n=4). Duringtherainy = +0.965, adults only: r = +0.952; n=4) and
season, decreasing abundanceof7". minorandB. decreasing soil temperature (P<0.05; adults and
browni(Fig. 3) was significantly correlated with nymphs: r = +0.967; n=4). During the rainy
increasing pH values at greater soil depths (T. season,verticaldistribution wassimilarbutmore
rmfoi=rn-oa0rd.:u9l5Pt1s<f0oon.rl0y5n;,yBmr.p=hb-rs0o.ow9nn7li0y:faPon<rd0a.dP0u<1l0t,.s0ra1=n,d-r0n=.y9-m90p8.h9fs9o,4r pfdorrsoocnmoor3up.ni5co-en7dscm(w,Feirg7.e%4t)a:fkr7eo3nm%f7ro-of1mt0ht.eh5e4ct8mo0pa(n3±d.529c7m%),pfs1re1ou%m-
adults and nymphs and r = -0.995 for nymphs thewhitesandsoillayer(10.5-14cm).About68%
only; n = 4, respectively). In M. tridens, the of the total catch were nymphs. No significant
increasing abundance of nymphs at greater soil correlation was found between the vertical dis-
depths was significantly correlated withdecreas-
tribution ofpseudoscorpions and changing con-
4i)n.gTshoieldmeocirsetausreeicnoanbtuenndta(nPc<e0.o0f5a,durl=ts-,0h.o9w77e;venr=, ditions ofsoil moisturecontent, soil temperature
and pH.
was significantly correlated with the decrease of
Seven species of pseudoscorpions were ob-
the soil temperature at greater soil depths
(P<0.05, r = +0.962; n = 4). This was also ob- tained during the dry and six species during the
+se0r.v9e5d1;fonr =ny4m)p.hsNeoofteTn.ymainndorpo(tPe<n0t.i0a5l, prar=- rreapionrytesdefarsoomnso(iFlisgo.fA5)m.azAolnliawnerderylparnedvifooruessltys
thenogenesiswereconfirmedforM. tridens,with (Mahnert and Adis, 1985). Chthoniidae were
sexually maturetritonyrnphs (= males)being ab- most abundant, with Tyrannochthonius (T.J
sent (cf. Adis and Mahnert, 1990a; Mahnert, minor Mahnert (dry season: 231 ±13 ind./nV\
1985). rainy season: 154±15 ind./m2), T. (T.) rotun-
dimanus Mahnert and Pseudochthonius
homodentatus Chamberlin accounting for 33-
PrimaryForeston White Sand Soil 45%ofthetotalcatch(Fig.5). Syarinidaewasthe
Pseudoscorpions accounted for 3-4% of all next most abundant family with Microblothrus
(ar1t4h,r0o0p0o-d1s5,ex0t0r0acitnedd./fmr2o,mA1cma"risoain1odfC1o4lclmemdbeopltah rtraiidneynsseMaashonn:er1t35(d±ry8sienads.on/:m2)16a4nd± I1d0eionbdi.s/imu2m,
db)i.srDeugrairndgedt)heindtrhyessetausdoyn,araebaou(tAd4is8%etoa/f.,th1e9t8o9taa,l srcahiunsytesreiasMoanhn14er±t4(dirnyd.s/ema2s)oanc:co2u9nt±i3ngifnodr./3m12-,
530(±22)pseudoscorpionswerecollected from 36% ofthe total catch. The Miratemnidae were
the top 3.5cm, 27% from 3.5-7cm, 12% from representedbyBrazilatemnusbrowniMuchmore
7-10.5cm (= humus layer) and 13% from the (17-35%; dry season: 91 ±6 ind./m2 rainy
,
white sandsoil layer(10.5-14cm). About66% of season: 168±16ind./m2).Lessabundant(^ 1%)
) ) 9
NEOTROPICALPSHUDOSCORPIONS AM
DRY SEASON RAINY SEASON DRY SEASON RAINY SEASON
N/m 14.4 L£ 9.6 14.4 N/m 154.052.9 1924.8 12S9 24.1
100 -i Ideobismm schusten 100 Tyrgnnocntornus minor
3N/m2=28.8 jN/m4-- 14.4 N/m" =230.9 2: N/m" = 154.0
50 - 50
Qa n
f—r
, ;
N/rr? 48.1 14 4 19.2 9.6 120.3 4.8 24.1 19.2 N/m 24.1 722 192 4ei 770 24,1 9.6 24.1
100 Brazilatemnus browni 'Ui Microbtotfrrus tndens
0) War = 91 3 iN/^ = 168 4 2 N/m2= 163-6 SN/m2= 134.
a
c
50 50
u
a>
a
Run 4-ATL A RaR
El
T r 1 r T I T
3.5 -70 -10.5 -44.0 -3.5 -70 -105 -14G -35-7.0 -10.5-140 -35 -7.0-10.5-14.0
Soil Depth (cm Soil Depth (cm
nymphs ] = adults
FIG. 6. Verticaldistributionoffourspeciesofpseudoscorpionsinsoil (%).Samplestakenevery3.5em todepth
of 14cmduringdry and rainy seasons in primary dryland forest on whitesand soil ncarManaus, Brazil.Total
catchofeachseason= 100%.Abundancegivenforeachsoillayer(N/m")andfortotalcatchperseason(XN/m2).
were Idcoroncidae [Albiorix gracilis Mahncrt) tween abundance and soil conditions was ob-
andChernetidae(Pseudopikmuscrassifemoratus served in T. minor: its abundance decreased at
Mahnert). The same four species (T. minor, B. greater soil depths as the soil moisture content
browni,M. tridensand/.schusteri)werethemost (P<0.05, r= +0.979 only foradults; P<0.10, r=
abundant in both seasons, however, their +0.935 for adults and nymphs, r = +0.900 for
dominance in the total catch per season varied nymphs only; n = 4, respectively) and the soil
(Fig. 5). In two species, vertical distribution dif- temperature decreased (P<0,05, r = +0.967 for
fered with the seasons. During the rainy season, adultsandnymphs, r-+0.972foradults; P<0.10,
0T.-7mcimnoarndanidn t/.hescthoupst3e.r5icwmeorfesofiolu,nrdesopnelctyivferloym. rN=o+s0i.gn9i3f9icfaonrtncyormrpehlsatoinolnyw;ans=f4o,unredspdeucrtiinvgelyt)h.e
Both species were found throughout the 0-14cm rainy season between the vertical distribution of
soil sample during the dry season (Fig. 6). Their speciesandtheabiotic factorsinvestigated. InA/.
tridens sexually mature tritonymphs (= males)
abundance was greatest in the top 3.5cm, inde-
were absent, which confirms neoteny and poten-
pendentofseasons. ThisisalsotrueforB. browni
which, together with M. tridens, occurred tial parthenogenesis in this species (of. 4.1.).
throughoutthe0-14cmsoiI sampleduring thedry DISCUSSION
season as well as the rainy season. Adults ofM,
tridenswere somewhat more abundantin the top
7cm. whereas no adults of/, schusteri were col- InCentral Amazoniatheabundanceofpseudo-
lected, probably due to low catch numbers. A. scorpions in primary and secondary dryland
arboricola was restricted to the top 3.5cm. forests on yellow latosol accounted (inde-
During thedry season, significantcorrelation be- pendently ofseasons) for3-5Tr ofall arthropods
438 MbMOIRSOFTHEQUEENSLANDMUSRIM
extracted from 1m* soil of i4cm depth and for ture in the primary forest on white sand mmI (cf,
10-15% when Ac-ah and Collembola were ex- 4.1,4.2.). Inaddition,decreasingabundancewith
cluded (cf. Adis e? aL. 1987a, b, unpublished depth was also related to somewhat lower
data). In the primary forest on white sand soil. temperatures recorded from the lower levels of
their abundance was only 0.7-0.9% and 3-4%, soil in Ihe white sand forest. During the rainy
respectively, due to largenumbersofPauropoda season,nocorrelationwasfoundbetweenspecies
and Diplura (cf. Adis & or*-. 1989a. b). About abundanceandsoil moisturecontentinthesetwo
two-thirdsofallpscudoscorpionsrecoveredfrom forest types. However, in the forest on yellow
the soil ofthe primary and secondary forest on latosol there was positive correlation between
yellow latosol (61-67% of (he total cjuhi and decreased abundance of nymphs and lower soil
about ihree-uUartcrs Of the primary white sand temperatures and negative correlation between
forest (75-84*1 inhabited the lop 7cm. No decreased abundance of nymphs, nymphs and
species were found to occur exclusively in the adults (P<0.05) andincreased soilpH. Ai greater
lower, mineral subsoil (e.g. in 10.5-14cmWsoWil soil depths there was an even higher statistical
depth). Two species (7". fniwrt H. bnwxn probability (P<0.01) of negative correlation be-
more abundant in the upper, organic layer tn the tween decreased abundance of adults and in-
three forests which were in tod iFigs 3, 6: creasedpH. Correlation between abundance and
fag.3inAdisand Mahncn. 1990a).M. tridens, T, il moisturecontenthasnotyetbeenreportedfor
- and B, hruymi were the most abundant T. minor from the secondary forest on yellow
species, representing89-95%ofthetotalcatchin latosol (Adis and Mahnert, 1990a). The more
theprimaryforestsonyellow latosolandonwhite homogeneousoccurrenceofA/,tridensin0-14cm
sand soil. They represented 5> f the total soil depth (Figs 3, 6; fig. 3 in Adis and Mahnert
catch in the secondary forest on yellow latosol, 1990a) wascorrelated with soil moisture content
where /. tenuis and A \ were frequent as as well: there was positive correlation between
well (22-33% of the total catch; cf. Adjs and decreasing numbers ofnymphsand sod moisture
Mahnert, 1990a). The similar spectuun ofpaeu- in the primary forest on yellow latosolduringthe
doscorpion species in forests on yellow latosol rainy season and, correspondingly, pegai
andon white sand soil andespecially the lack ot relation between increasing abundance of
endemic species in the latterforest type, confirm nymphs and decreasing soil moisture content at
the geological results ufChauvel fll aL (1987). greater soil depths in the secondary forest on
They reported that the white sand area inves- yellow latosol (cf Adis and Mahnert, 1990a) In
tigatedrepresentsthefinal stageofpodzolisation, adults, howeveT thcic wn% a posiiive correlation
i.e. the transformation ofclayey latOSOla to white between decreased abundance il moisture
sandy podzols by (long-term) Weathering And content (secondary forest) and between
leaching processes [fthewhitejtmdswc decreased abundance and lower temperatures at
large dried-upold riverbed, species composition greatersoil depths (primary
would be different Irom the yellow latosol area Thett data all suggest if ips other than
id Adi* aid Mahncn 1990b). soil moisture content (abiotic and/or bioticones)
Neitherduring thedry northerainy season: js welt cause the different distribution of
Ihe abundance of pscudoscorpions in mineral pseudoscorpion species in the soil. A significant
subsoilshigherinresponsetothechangingmois- correlation, forexample, was found between Ihe
ture content in organic layers. Thisphenomenon ahun<LoKe of pseudoscotpions and Collembola
has been ^ported in arthropods in the seasonal- in thesecondaryforestonyellow latosol (P<0 05;
ly-dry tropical forests, where periods without dry season r = +0,969,rainy season: r= +0.989;
precipitation occur |cf. climate diagrams for n=4) andinthepiimary forestonwhite sandsoil
CentralAmerica(WalterandLietli. I960 67)and (dry season: P<0.01, r = +0.991, rainy sc,
for Manaus (Worhcs, I986)J. Higher or lower P<6.05. r= +0.967; n - 4). This relationship hiiN
abundanceof(at least some)speciesat adistinct to be investigated in more detail, as sprii.^tail-
soil depth seems lo he related todifferent abiotic representapotential food source forpscudoscor-
factors. For example, the decreasing abundance pions (cf. Jones, 1975; Rcssl and Bcier, 1958;
of71 minorwith soildepthduring the dry season Wcygoldt, 1969). Thus, complementary studies
correlated withsoil moisture content in two should be carried out on reproduction patterns,
fbfl&l types: negatively with in E 'Moisture food availability and trophic structure over a
content values in the primary forest on yellow one-yearperiod (cf. Adis et aL, 1988). Data also
latosol and positively withdecreasing soil mois- show ihat the number of samples taken per soil
1
NEOTROPICALPSEUDOSCORPIONS
•1.i-j
layer (n= 6) was sufficient for a species Verticaldistributionandabundanceofarthropod*
dominance of5% ofthe total catch, as standard inthesoilofaNeotropicalsecondaryforestduring
deviation did not exceed 203?. Results indicate diedrvseason.TropicalEcology 28: 374-1S
thatterncolepseudoscorpionsofprimarydryland ADIS.J.. MORAIS.J.W.DE& MESQUITA.H.G.DE
forests on white sand soil and of secondary 1987b. Vertical distribution and abundance of
arthropods in thesoilofaNeotropical secondary
dryland forests on yellow latosol represent the
forest during the rainy season. Studies on
lerricole species spectrum ofa primary dryland NeotropicalFaunaandEnvironment22; 189-197
forest on yellow latosol, which has been altered ADIS, J., MAHNERT, V.. MORAIS, J.W.DE <fc
titherby geomorphological/geochermcalproces- RODRIGUES, J.M.G. 1988. Adaptation of an
ses or human impact (ef Morals, 1985; Amazonian pseudoscorpion (Arachnida) from
Rodrigues, 1986). Habitat preference in pseudo- drylandforeststo inundation forests. Ecolocv 69
scorpions and the many habitat types existing in 287-29].
Amazoniaareconsidered tobe majorreasonsfor ADIS. J . MORAIS. J.W.DE, RIBEIRO, E.F. 4:
the high species diversity found in neotropical RIBEIRO, J.C. 1989a. Vertical distribution and
pscudoscorpions (cf. Adis and Mahnert, 1985, aabNuenodtraonpciecaolfcaarmthprionpaordasnafrfoormeswthidtureisnagntdhesorialinoyf
1990b).
season Studies on Neotropical Fauna and En-
vironment24: 193-206.
ACKNOWLEDGEMENTS
ADIS. J., RIBEIRO. L K. MORAIS, J.W.O
CAVALCANTE,E.T.S. 1989b.Verticaldistrihu
The pseudoscorpion material was obtained lionandabundancenfarthropodsfromwhitesand
dFiuerlidngEctohleopgoys*t-ogfraIdNuPaAt/eUcnoiuvresressit*EynAimoamzoolno£asicaatI dsoriylosfeaasNoeno.trSotpuidciaelscaomnpiNneaortarnoapifcoarlestFdauurniangntoltt!e
Manaus (Brazil) in August 1988 and Tico-en- hrivironment24; 201-21 1,
tomologicalcourseonneotropical arthropods' of ANDERSON, A.B. 1981. White-sand vegetation ol
BrazilianAmazonia. Biotropiea 13: 199-210.
the Chrisiian-Alhrechis-UniveTsity at Kiel (Ger- ANDERSON, A.B., PRANCE, GT. & ALBUQUKR
many) in October 1990. All participants are QUE.B.w.P.DE i975.£srudoesdtweavege
thanked for their valuable collaboration. Inge dasCampinas Amazotucas -III. A vegetacao Jen-
Schmidt-Brockmann kindly typed the hosa da Campina da Reserva Biologica INPA
manuscript, Susannc Hamann (both at MP1 in SUFRAMA (Manaus-Caraearaf, km 62). Acta
madethe drawingsand Nancy Wetdercor- Amazonit -246.
rectedthe Englishdraft. BECfc L, [964, Tropisehe BodcitftuitB im Wecbsrl
von Regen-undTrocker^ek Natur und Museum
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