AMAZONIANA x 297 -314 Kiel, Dezember 1985

From cooperation between Max-Planck-Institute for Limnology, rvVorking group "Tropical Ecology",

pttin, W"tt Germany, and Instituto Nacional de Pesquisas da Amazõnia, Manaus-Amazonas, Brazil

Da cooperação entre Max-Planck-Institut für Limnologie, Arbeitsgruppe Tropenökologie, Plön,

Alemànha Oc., e Instituto Nacional de Pesquisas da Amazônia, Manaus-Amazonas, Brasil

on the natural history and ecology of Pseudoscofpiones (Arachnida)

from an Amazonian blackwater inundation forest*

by

Joachim Adis and Volker Mahnert

Dr. Joachim Adis, Max-Planck-Institute for Limnology, Working Group "Tropical Ecology"'

PO-Box 165, D - 2320 Plön, F.R.G.

Dr. Volker Mahnert, Museum d'histoire naturelle, Cæe Postale 434, CH ' 121 1 Genève 6,

Switzerland(accepted for publication: May 1985)

AbstractThe habitats and changes in habitat of 20 pseudoscorpion species in response to up to six months

forest ínundation are presented. The fauna is divided into 16 nonmigrating and 4 migrating species (2

terricolous and2æboicolousspecies).Adaptationsof life-cycletoforestfloodingandseasonaltreetrunk migration are discussed for Tyrannochthonius amazonicus, T. rnigans (Chthoniidae), Braiiløtemnus

browní (Mftatemnidae) and Pachyolpíum irmgardae (Olpiidae) based on data from trunk traps (arboreal

photo-eclectots), emergence traps on the forest floor (ground photo-eclectors), soil extraction (I(EMPSON

methotl), canopy fogging (pyrethrum method) and the collection of epiphytes.

Kêywords: Pseudoscorpiones, soil fauna, inundation forests, Neotropics, Brazil'

Resumo

Os habitats e mudanças de habitat de 20 espécies de Pseudoscorpiones são apresentados em

relação à até seis meses de inundação de floresta. A fauna é dividida em 16 espécies não-migrantes e 4

espécies migrantes, sendo 2 terricolas e 2 arborícolas. Para Tyrannochthonius nmazonicas, T. migrans(Chthoniidae), Brazilatemnus browní Mfualemnidae) e Pachyolpium ínngardae (Olpiidae) são discutidas

as adaptações de ciclo de viila à inundação de floresta e migração sazonal nos troncos de âwores, baseadas

em dados de armadilhas de tronco (foto-eclectores de árvore), armadilhas de emergência no chão da

floresta(foto-eclectores de solo), extração de solo (método de KEMPSON), nebulização de copa de

áwores (método do piretro) e a coleção de epífitas.

* This study is dedicated to Prof. Dr. Harald Sioli to commemorate his 75th birthday

0065-6755 | 1985 I 29'7 I @ MPI für Limnologie, AG Tropenökologie' Plön; INPA, Manaus

291

1. Introduction

Riven in the Central Amazon show annüal water-level fluctuations which may be asmuch as 14 m between high and low water (Rio Negro at Manaus). consequently vætriparian forest areas representing abolt 2 % of ¡he Brazlian Amaz on region, are floodedfor several months of the year. A periodically inundated forest, flooded by regular annualcycles of blackwater rivers is termed seasonal igapó @RAN cE 1979). rn 1976 - 7 jarthropod reaction to inundation was studied as part of a "minimal program for ecosystemanalyses" in the Manaus area (ADIS 1981). Activity density (,,Aktivitätsdichte";SCHWERDTFEGER 1975) of arthropods on the forest floor was detected using groundphotoeclectors and pitfall traps. Arboreal photo-eclectors, mounted on tree trunks,monitored trunk ascent and descent by arthropods. The non-flying and limited-flyingarthropod faunæ ofthe seasonal igapó were classifìed into terricolous and arboricolousgroups' They consisted of a) nonmigrants, wtrich chiefly remained either in the trunk andcanopy region, or on the forest floor and survived forest inundation; b) migrants, whicheither moved onto tree trunks and into the canopy before flooding and/or descended tothe forest floor during the non-inundation period (ADIS lgsl; ADIS & SCHELLER 19g4;SCHELLER&ADrS 1984).

Eigþteen species ofPseudoscorpiones were reported from the seasonal igapó understudy (ADIS 1981; MAHNERT 1979). Out of these, eleven species were considered barkinhabitants of tree trunks, representing the arboricolous nonmigrating group. Six specieswere considered litter inhabitants, representing the terricolous migrating group whichsurvived the 5 - 6 month inundation period in the trunk/canopy area. However catchesfrom ground photo-eclectors and forest floor pitfall traps didïot contain the earliestdevelopmental stages (proto- and deutonymphs). Thus, the life cycle of terricolousPseudoscorpiones, bæed chiefly on trunk captures, remained to be confìrmed. For thisreason, pseudoscorpion distribution in the soil of the seasonal igapó was investigated inl98l ' 1982 at the same study site as that used in 1976 - lg77. Furthermore, occurrenceofPseudoscorpiones in tree crowns was studied by canopy-fogging and the collection ofepiphytes (cf. MAHNERT 1985a).

298

2. Study area and methods

The study site was situated on the lower course of the Rio Tarumã Mirím (03" 02'S, 60. 1 7'W),a tributaly of the Rio Negro, about 20 km upstream of Manaus. The seasonal þapó possessed buttressedfiees up to 35 m high \¡¡ith an almost closed canopy, colonized by numerous epiphytes but had fewherbs and no shrubs. Forty-seven tree species at densities between 1,300 and z,gbo trees/tra were recordedin the study area. Ten species had tiunk ci¡cumferences of ), 100 cm of which Atilinø lafifona BENTHvar.løtifolía (Leguminosae) was dominant, The clayish-sandy sof was covered by a 5 - l0 cm thick finehumus layer which was penetrated by a matting of¡oots, A detailed description and characteiization ofthe study area, as well as information on the classification of inundation foìest types, is given by ADIS(1981, 1984), IRION &ADrs (1979), TRMLER (1975, 1977) and pRANcE oítb-t.rn"igapó understudy was submerged under floodwater, up to 3.35 m from March/April to August/Septembår. It wassubject to a rainy season (December-May) and a tlry season (June - Nàvember; RIBEIRO & ADIS 19g4).Distribution of Pseudoscorpiones in the soil was studied between September i981 an¿ February l9g2(non-inundation period). Once a month six L4 cm deep soil samples were taken at random intervals along

alràîsect using a split corer (= steel cylinder with lateral hinges; diameter 21 cm) driven into the soü

by a mallet, and then divided into four subsamples of 3.5 cm each. Animals were extracted from sub-

samples following the method of KEMPSON et al. (1963). Pseudoscorpiones we¡e collected from tree

øowns by fogging canopies with pyrethrum in the early dry season (July 1977, 19'19), when the sea-

sonaligapí was completely flooded (ADIS et aL 1984; ERWIN 1983). In addition Pseudoscorpiones

were sampled from bromeliads (Aechmea setigeraMART. ex SCHULT. F., Streptocalyx poeppigii

BEER) 5 - 25 m above ground in June 1981 and August t979 (earLy dry season, igapó flooded) as well

as in October 1980 (dry season, igapô not flooded). - AllPseudoscorpiones collected were classified

as juveniles (protonymphs, deutonymphs, tritonymphs) or adults (males and females).Species are listed in alphabetical order by families. The taxonimic work for this paper was done

by V. Malnert, the collection and evaluation of field data by J. Adis.

3. Results and Discussion

A total of 696 Pseudoscorpiones tr¡r'ere extracted from soil samples. The majority(89%of t}re totalcatch)werejuveniles: lTTprotonymphs (25Vù,193 deutonymphs(28 Vù and 250 tritonymphs (36 %).There was a I : 2 sex ratio of adult males and females.

About 83 % of all specimens were caught between 0 - 7 cm depth, irrespective of theirdevelopmental stage. The higþest population density, of 1,020 ind. m-2, wÍIs recorded inJanuary 1982 (0 - 14 cm depth) and the lowest, of 183 ind. m-2, in September 1981, thefirst month of the non-inundation period. Three species were collected from seasonaligapó soils: Tyrønnochthonius umazon¡cøs MAHNERT,T. migrøL,ts MAHNERT andBrazilatemnus browni MUCHMORE. Eleven species were obtained from epiphytes andduring canopy-fogging (Table 1).

Catch data, together with previous capture data from pitfall traps, ground andarboreal photoeclectors (ADIS 1981) and various standing crop samples, provided con-clusive information on the life cycle, habitat and ecology of pseudoscorpion species fromthe seasonal igapó.

299

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(lnd.)n=319

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ffi inundotion period f-l non - inundotron period

P = protonymphs, o = deutonymphs, r= tritonymphs, w= femoles.M= moles

Fig. l:Activity density of Tyrannonchthonius amozonicus (Chthoniitlae) on the forest floor (1 _ g groundphoto-eclectors (E; Ind. m-z), 2 pi$all traps (BoF), trunk ascents and trunk descents (3 arborealphoto-eclectors respectively (BEî , BE J )) betweerr January l9'7 6 and May 1977 at Rio TarumãMirim. (Data from ADIS 1981, modified).

3,5-7,0 10,5-1 4,0

Figs. 2 - 4:2: Temporal occuÍence of developmental stages of Tyrønnochthoníus amazonic¿ls (Chthoniidae) inthe soil (0 - 14 cm depth). Monthly samples taken during the non-inundation period (September 1981 -

February 1982) af Rio Tarumã Mirím,3: Distribution of Tyrønnochthoníus amazonícøs (Chthoniidae) in the sotl, l%). Samples taken every

3.5 cm to a depth of 14 cm between September 1981 and February 1982 (non-inundation period) at

Rio Tarumã Mirím; total catch = 100 %.

4: Percentage of developmental stages of Tyrannochthonîus amazonicus (Chthoniidae) caught in thesoil (0 - 14 cm depth) bet\ileen September 1981 and February 1982 (non-inundation period) at RioTarumã Mirím; total catch = 100 %.

no co UTES

302 303

T. amøzonicus is considered univoltine. Reproduction occurred on the forest floorand was well synchronized with the non-inundation period. In the soil development fromegg to tritonymph took a maximum of four months. The species is a representative of theterricolous migrating arthropod group (ADIS 1981) adapted to annual flooding.

Chthoniidae : Ty ra nno c h tho niu s migra ns MAHNERTT' migrans ís the second noctumal species considered endemic to the seasonal igapó.

As in the closely related T. amazonicus, migratory tritonymphs survived forest inundation(April - August) in the trunk/(canopy) region. As the water ieceded they recoloni zed. theforest floor for moulting and subsequent reproduction (Fig. 5, BE.t). However it remainsto be shown whether tritonymphs and/or adults have a period of aátivation in the soilthroughout the dry season before this (september - November). The progeny (= proto-nymphs) were only observed from December onwards, the beginning-of the rainy season(Fig. 6)' In contrast ro T. amazonicus, T. migrøns inhabited the lower humus layer. Eighty-fow % of all juveniles and adults were caugþt between 3.5 - 10.5 cm depth (rig. z). rJpuia-tion density of T. migrans in the soil was^only half that of T. amazomcøs. Highest nurnãr6were observed in December, 2g4 ind. m-2 for juveniles and adults, 96 rr,d.

^i2 fo, p.otÀ_

-

nymphs and 9l ind. m-2 for tritonymphs (0 - 14 cm depth; Fig. 6). No difference wasfound between the total numbers of specimens caught fer ny*pt ut stage during the non_inundation period (Fig. 8). Protonymph to tritonymptrdeveiopment was fasterln T. migransthan in T. amazonians (Figs. 2,6), ptobably being concluded within 4 weeks. Highest trito-nymph activity density in ground photo-eclectors was recorded in February, one monthearlier than for T. amuzonicus (Figs. 1, E; 5, E).

As in Z amøzonicus, th'e number of tritonymphs in the trunk region was considerablyreduced by predatory arthropods during forest inundation. No specimãns of r. migrønswere collected in the upper tree crown with canopy-fogging, from epiphytes or duringstanding crop sampling, Reproduction occurred on t¡e forest floor, T. migrans thus belongsto the terricolous migrating arthropod group (ADIS lggl).

closely related to T. amazonicus, the univoltine T. migrøns is separated from itspotential competitor by spatial, temporal and onthogenetic differencr, ¡"r. wooo aGABBUTT l97S).

Miratemnida e: Brazilatemnus browniMUCHMOREB' browni is a frequent noctumar species in the Amazon region, inhabiting dryrand

forests, seasonal igapó forests and blackwater swamp forests, which have permanentlywaterlogged ground (MAHNERT 1979; MUCHMORE 1975).In primary and secondarydryland forests near Manaus, B. browni was only found in ttre soii and never caught ontree trunks or in the canopy. There was no distinct reproduction period and proionymphswere found throughout the year (ADIS et al. 1985; Á¡ts ¿. scH'uBART 1gg5). In theseasonal igap6, B. brownihad two reproductive periods: a major on. in th"iruít¡t.*pylregion during forest inundation (April - August) and a second, minor one, on the forestfloor during the non-inundation period. About'three times as'many unirnutr, predominantlytritonymphs, were caught during trunk descents (Fig. 9, BE+)

^ ,årp.rr¿ to trunk ascents(Fig. 9, BEl). After recolon2ing the forest floor, tritonymphs moulted and reproduced

(Fig. 10). Only a few animals, exclusively adults, were caught within the first two monthsof the dry season (i. e. September/October). As in Z. migrans the cause has yet to be

determined. First free-living protonymphs of the new generation were recorded in Novem-

ber and first tritonymphs in January Gig. 10). Brood or moulting chambers (GABBUTT &VACHON 1967; WOOD & GABBUTT 1979a, b) were not observed. Eigþty-six % ofjuveniles and adults were extracted from soil samples during the rainy season (December -

February). Their population density was highest in December with 390 ind. m'2 (0 - 14 cmdepth).

The highest number of tritonymphs was recorded in January with 24I ind. m-2 @ig.lO). B. browni was mostly recovered from the lower humus layer with '77 % of all speci-

mens caught between 3.5 - 10.5 cm depth (Fig. 1l). More tritonymphs were collectedduring the non-inundation period than proto- and deutonymphs (Fig. 12). Abundance ofadults was low and we assume that the parent generation had died by the beginning of theinundation period (March/April). B. browni was scarcely detected on the soil surface. Fewanimals were caught in ground photeeclectors shortly before forest inundation (Fig. 9, E)and none in pitfall traps. [n contrast to T. qmazonlcus and T. mígrans in which trunkascents were initiated at the beginning of the rainy season (December), B. browni oríycommenced migration a short time before the forest floor was inundated (Fig. 9, BEl). In1976 rhe highest capture rates in arboreal photo-eclectors were recorded one week beforeforest inundation. The high wetness and relative humidity on the forest floor, caused byrising waters, apparently forced all developmental stages into the trunk area (Fig. 9, BEl;cf. Fig. 14 in ADIS 1981). Some animals must have moved up the trunk with rising water-level, as specimens were caught until June, when flooding had reached its maximum (Fig.9, BEl). We presume that tritonymphs moulted to adults in the trunk area where the mainrcproductive period occurred in April/May. Tritonymphs of the new generation were cap-tured during trunk descents, together with some remaining proto- and deutonymphs, fromearly July onwards, i. e. six weeks before the start of the non-inundation period (Fig. 9,BE.l). Heavy leaf-fall combined with strong insolation in July/August (ADIS et al.1979)may have initiated early trunk migration. B. browni wæ not found in epiphytes. Trito-nymphs were obtained in low numbers with canopy fogging during forest inundation (inJuly).

B. browni is considered bivoltine. Originating from adjacent dryland forest soils,this species seems to have colonized and adapted to the seasonal igapô secondarily. Proto-nymph to tritonymph development læted a maximum of three months and was sosynchronized with water-level fluctuations that migrating tritonymphs stages were attainednear the beginning and the end of the non-inundation period. Main reproduction occurredin the trunk region during forest inundation.,B. browni thus represents the arboricolousmigrating arthropod group (ADIS 1981). The species is separated from Z amazonicus andT. migrans by spatial, temporal and onthogenetic differences (cf. wooD & GABBUTT1978).

305304

Fig.5rorny seoson > dry seoson ---> ro¡ny seoson ---+

BEl 1900

1 600

1 300

1 000

700

400

100

50

0

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r00

400

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Tvronnochthonius mi qro ns

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ffiinundotion per¡od f--l non-inundotion period

e= protonymphs, o=deutonymphs,r=tri tonymphs, ui= f emoles, M : moles

Fig. 5:Activity density of T¡rannochthontus migrans (Chthonüdae) on the forest floor (1 - 5 ground photo-eclectors @; Ind. m-¿)), trunk ascents and trunk descents (3 arboreal photo-eclectors respectively(8E1, BEJ)) between January 19'16 and May 1917 at Rio Tarumã Mirím.(Data from ADIS 1981, modified).

306

20 (lnd.) Fig.6Ty ronnochthonius miqrons

n =104

10

s o N D F

1980 1 981

ffiprotonymphs fldeutonymphs ffitritonymphs Qfemoles ! motes

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3,5-7,0

Figs. 6 - 8:6: Temporal occuÍence of developmental stages of Tyrannochthoníus migTans (Chthoniidae) in thesoil (0: 14 cm depth). Monthly samples taken during the non-inundation períod (September l98l -

February L982) at Rio Tarumã Mirím.7: Distribution of Tyrannochthonius migrans (Chthoniidae) in the soil (%). Samples taken every 3.5cm to a depth of 14 cm between September 1981 and February 1982 (non-inundation period) at RioTarumã Mirím; total catch = 100 7o.

8: Percentage of developmental stages of Tyrannochthonius migrøns (Chthoniidae) caught in the soil(0 - 14 cm depth) between September 1981 and February 1982 (non-inundation period) at RioTarrimã Mirím; total catch = I00 %.

A(16,3

307

Fi g.9<- fo(lnd )

ny seoson ry seoson rorny seoson11 00

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100

50

0

50

100

Brozilotemnus browni n=2.3!-2

trunk oscents

./D_D

wP 0

-l-T-T

600

1100

1 600

2100

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31 00

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8E,r 4100

4600

830

irunk descents n=6,235

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15

1976 1977

ffi inundotion period l---l non _ inundotion periodp

= protonymphs, o= deu tonymphs, r= tri tonymphs, w = femoles

Fíg. 9:Activity density of Brøziløtemnus browni (Miratemnidae) on the forest floor (1 - 5 ground photo-99lectorl (E; Ind. m-2)), trunk ascents and trunk descents (3 arboreal photo-eclectors respectively(8E1, BEI) between January 19'16 andMay 1977 at Rio Tarumã Mi¡-ím.(Data from ADIS 1981, modified).

res

308

60 (ln d. )Fis. 10

Broziloiemnus browni

n=22t

LO

20

S 0 N D F

r980 1981+I molesfr protonymphs fl deutonymphs ffi tritonymphs Qnemoles

80 t%) Fig.11 Fig.12n =22t,

ffiffi

JU ve n le

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0-3,5 3,5-70 Z0-10,5 10,5-14,0 cmP = protonymphs,D = deutonymphsT = tritonymphs, A = odutts

Figs. l0- 12:10: Temporal occurrence of developmental stages of Brazilatemnus browni (Miratemnidae) in the

soil (0 - 14 cm depth). Monthly samples taken during the non-inundation period (September 1981 -

February 1982) at Rio Tarumã Mirím,tl.: Distribution of BrøzíIøtemnus browní (Miratemnidae) in the soil (7o). Samples taken every 3.5 cm

to a depth of 14 cm between September 1981 and February 1982 (non-inundation period) at RioTarumã Mirím; total catch = 100 %.

12: Percentage of developmental stages of Brøziløtemnus browní (Miratemnidae) caught in the soil(0 - 14 cm depth) between September 1981 and February 1982 (non-inundation period) at RioTarumã Mirím; total catch = 1,00 %,

132,6%lT

309

3.2. Nonmigtating Pseudoscorpiones

Sixteen species which are believed to live under loose bark on trunks and/or in epi-

.tvtes of trees were collected in the seasonal igapô. Some of them occurred occasionally

ãnîr roif surface during the rainy season and were caught in ground photo-eclectors

üOfS f SSf ). h many cæes, they may have been washed down the tree trunks by heavy

ìJnr. Uort species were caught in low numbers and only preliminary statements on their

talunlhistory and ecolory are possible at this stage'

Atemnidae : Paratemnus minor (BALZ/JI{\This species was collected under loose bark of trunks to a height of 20 m, in arboreal

ohoto-ecleciors (ADIS 1981), but not in epiphytes. Nymphal instars occurred througþout

ih. yrut and constructed moulting chambers. Females with eggs were found in brood

.fr*trtr in October and March. One female was recorded in ground photo-eclectors (in

December).

cheme tidae : A m erich em e s b e thaniae MAHNE RT / A. inc ertu s MAHNE RT

All developmental stages of both species were caught in arboreal photo-eclectors.

Somewhat higþer capture rates wele recorded fot A' bethaníøe duingthe dry season (cf.

Table 8 in ADIS 1981). Females of/. incertus were also obtained by canopy'fogging and

one male wæ recorded in ground photoeclectors (in September).

Chemetidae : C erio cherne s amaz onicu s MAHNERTThis species was only found in epiphytes between 13 - 19 m above ground. All

developmental stages were present (Table 1).

Chemetidae : Lustrochemes intermedius (BALZAN)

only one female was obtained during canopy-fogging in July 1979 (cf. MAHNERT

1e8sb).

Chemetidae : Pachy chernes baileyí FEIOAll developmental stages were caugþt throughout the year in arboreal photo'eclectors

(ADIS 1981). The species was also collected by canopy-fogging, from epiphytes at aheight

of l3 m (Table 1) and under loose bark of Aldina latífoliø (Leg.) up to 20 m above ground.

There was no evidence that brood and moulting chambers are constructed. Higher captule

rates were recorded during the dry season.

cherne tidae : P arach erne s meinertii (MTH) /P. melanopy gu s BEIER /P. plumo su s

(WITH)/P. setiser MAHNERTThese four species occurred in very low numbers in arboreal photo-eclectors (ADIS

1981). Of P. meinertü andP. setiger only adult instars were caught while deuto- and trito-nymphs of P. plumows were also collected. Protonymphs and a female with eggs of P.

melønopygus were found under loose barkT - 10 m above ground onAldina latiþlia (Leg.)

trunks during forest inundation (in July).

311

Garypidae : G eogary pu s amøz o nicu s MAHNERTG. ømazonicus wæ initially conside¡ed a terricolous migrating species (ADIS 19gl).

Its absence on the forest floor indicates, however, that the species belongs to the arbori-colous nonmigrating group. G. amøzonícus was caught throughout the year in arborealphotoeclectors. Higher capture rates were recorded during the dry season. protonymphsoccurred between July and March. Tritonymphs and adults were also found in epiphyìesin 13 - 25 m height (June, August, October).

Tridenchthonüdae : Tridenchthoníus brqsíliensis MAHNERTonly one female was caught in an arboreal photoeclector (ADIS 19gr).

withiidae: Dolichowithius intermedius MAFINERT/ D. mediofascørrzs MAHNERT/D. minutus MAHNERT

These three species occurred in low numbers in arboreal photo-eclecton (ADIS 19g1).D. intermedius and D. mediofascíatur were also obtained with canopy fogging (Table l).All developmental stages were caught only in D. mediofasciatus. rts ptofonymphs wereobserved between september and December and the species or.urrrã somewhat morefrequently ín the lower trunk region during the dry seãson.

wi thiidae : P ara wi th iu s (vi c t o r wi th iu s) gr ac í r i m anus MAHNERT

. Low numbers of protonymphs, deutonymphs and adurts of this species were caughtduring the dry season in arboreal photo-eclector, qalts teAt;.

¡ nr( J. & U. SCHELLER (1984): On the natural history and ecology of Hanseniella arborea (Myriapoda,

^"'"'Sy.phyla, Scutigerellidae), a migrating symphylan from an Amazonian black-water inundation

forest.- Pedobiologia2T (1): 35 - 41'

^ nrc r & H. SCHUBART (1985): Ecological research on arthropods in Centrai Amazonian forest-o"'"' å.orutpms, with recommendations for study procedures.- In: COOLEY, J.H. & F.B. GOLLEY

(eds.): Trends in Ecological Research for the 1980's: 111 - 114, NATO Conference Series:

Ecology, Plenum Press, New York' 344 pp.

aDtS. J., MAHNERT, V., MORAIS, J.W. DE & J.M.G. RODRIGUES (1985): Adaptations of the

pseudoscorpion Brazilatemnus browni MUCHMORE (Miratemnidae, Pseudoscorpiones,

Arachnida) from Central Amazonian dryland forests to blackwater inundation forests.- (in prep.).

ERWIN, T.L. (1983): Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled

by insecticidal fogging.- In: SUTTON, S.L., WHITMORE, T.C. & A.C. CHADWICK (eds.):

Tropical Rain Forest: Ecology and Management.- Proc. Trop. Rain Forest Symp. Leeds 1982:

59 - 75. - Blackwell, ScientificPubl', Oxford.

6ABBUTT, P. D. (196 7) : Quantitative sampling of the pseudoscorpion Chthonius ischnocheles ftombeech litter.- J . Zoo1., Lond. 15 1: 469 - 47 8.

GABBUTT, P.D. (1969): Life histories of some British pseudoscorpions inhabiting ieaf-1itter.- InrSHEALS, J.G. (ed.): The Soil Ecosystem: 229 - 235. Systematics Association Publication No. 8,

London.GABBUTT, P.D. (1970): Sampling problems and the validity of life history analyses of pseudoscorpions.-

J. nat. Hist. 4: 1 - 15'

GABBUTT, P, D. & M. VACHON (196 3) : The external morphology and life history of the pseudo-

scorpion Chthonius ischnocheles (Hermann).- Proc. zooi. Soc. Lond. 140 (1): 75 - 98,

IRION, G. & J. ADIS (1979): Evoiução de flo¡estas amazônicas inundadas, de igapó - um exemplo do

rio Ta¡umâ Mirím,- Acta Amazonica 9 (2): 299 - 303.IRMLER, U. (1975): Ecological Studies of the Aquatic Soil Invertebrates in Th¡ee Inundation Forests

of Central Amazonia.- Amazoniana 5 (3): 337 - 409.

IRMLER, U. (19'1'l): Inundation-forest types in the vicinity of Ma¡aus.- Biogeographica 8t 17 - 29.

IGMPSON, D., LLOYD, M. & R. GHELARDI (1963): A new extractor of woodland litter.- Pedobio-

logia3l I - 21.MAHNERT, V. (1979): Pseudoskorpione (Arachnida) aus dem Amazonasgebiet (Brasilien).- Revue

suisse Zoo1. 86 (3): 719 - 810.

MAHNERT, V, (1985a): Weitere Pseudoskorpione (Arachnida) aus dem zentÍalen Amazonasgebiet(Brasilien).- Amazoniana 9 (2): in press.

MAHNERT, V. (1985b): Pseudoscorpiones (Arachnida) from the Lower Amazon region.- Revista Bras,

Ent., in press.

MORAIS, J. W. DE (1985): Abundância de distribuição vertical de artrópodos do solo de florestaprimária não inundada.- M. Sc, thesis, INPA/Manaus (Brazil), 95 pp.

MUCHMORE, W. B. (1975): Two miratemnid pseudoscorpions from the Western hemisphere (Pseudo-

scorpionida, Miratemnidae).- The Southwestern Naturalist 20 (2); 231 - 239,PRANCE, G,T. (1979): Notes on the Vegetation of Amazonia, III. The terminology of Amazonian

forest types subject to inundation.- Brittonia 3l (1): 26 - 38,RIBEIRO, M. DE N.G. & J. ADIS (1984): Local rainfall - a bias for bioecological studies in the

Central Amazon.- Acta Amazonica, in press.RODRIGUES,J.M.G,(1985): Abundânçiaedistribuiçãoaoverticaldeartrópodosnosolo-de

capoeira - em região de água preta.- M. Sc. thesis INPA/Manaus (Brazil): in preparation.SCHELLER, U. & J. ADIS (1984): A new species of Ribautiella (Myriapoda, Symphyla, Scolopendrelli-

dae) from an Amazonian black-water inundation forest and notes on its natwal history andecology.- Amazoniana 8 (3): 299 - 310,

SCHWERDTFEGER, F., (ed.) (1975): Synökologie. Paul Parey, Hamburg, Berlin,45l pp.WOOD, P.A. & P,D. GABBUTT (19?8): Seasonal verticai distribution of pseudoscorpions in beech

litfer.- Bull. Br, arachnol. Soc.4 (4): 176 - 183.

313

4. Acknowledgments

Special thanks a¡e due to IRMGARD ADIS for sorting thousands of pseudoscorpiones and toour technical staff, especially VERA BOGEN and EDILSON ARAúJO DA SILVA, fbr the enormouseffort in the field and laboratory. Dr. E.J. cox tMpI, plön) and D¡. p.D. GABBUTT (university ofManchester) are cordially thanked fo¡ valuable discussion ald suggestions regarding manuscript format.ELKE HARMS (MPI, Prön) kindly made the drawings, v. KRANTZ (r.{at. Museum Nat. History(smithsonian Institution)) took the photograph of the brood chamber (Fig. 13). Dr. WLLIAMRODRIGUES, INPA, Manaus, kindly identified the plant material.

5. References

ADIS, J' (1981): Comparative ecological Studies of the terrestrial arthropod fauna in CentralAmazonian Inundation-Forests.- Amazoni anaT (2): g7 _ 173.

ADIS' J' (1984): "Seasonal igapó"-forests ofCentral Amazonian blackwater rivers and theirterrestlial arthropod fauna.- In: sIoLI, H. (ed.): The Amazon - Limnology and landscapeecology of a mighty tropical river and its basin: 24s - 26g. Monographiaeîiorogicae, JunkPubl., Dordrecht, Boston, Lancaster, 763 pp.

ADIS, J'' FURCH, K. & U. IRMLER (1919): Liiter production of a Central Amazonianblackwaterinundation-forest.- Trop. Ecol. 20 (2): n6 _ t4S.

ADIS, J., LUBIN, Y.D. & G.G. MoNTGoMERy (1984): Arthropods from the canopy of inundatedand ter¡a firme forests near Manaus, Brazil, with critical cånsiderations on the pyrethrum-foggingtechnique.- Stud. Néotrop. Fauna & Envi¡on. l9 (4): 223 _ 236,

312