On the Abundance and Phenology of Palpigradi (Arachnida) from Central Amazonian Upland Forests

7 Pages·1997·6.7 MB·English

by  J Adis

1997. The Journal of Arachnology 25:326-332
ON THE ABUNDANCE AND PHENOLOGY OF
PALPIGRADI (ARACHNIDA)
FROM CENTRAL AMAZONIAN UPLAND FORESTS
J. Adis’, U. Scheller^, J.W. de Morais^ B. Conde'’ and J.M.G. Rodrigues^: ’ Max^
Planck-Institute for Limnology, Tropical Ecology Working Group, Postfach 165, D-
24302 Plon, Germany. ^Haggeboholm, Haggesled, S-53194 Jarpas, Sweden. ^Instituto
Nacional de Pesquisas da Amazonia (INPA), C.R 478, 69.011-970 Manaus/AM,
Brazil. '’Musee de Zoologie de TUniversite et de la Ville de Nancy, 34, rue Sainte-
Catherine, F-54000 Nancy, France.
ABSTRACT. The 745 palpigrads (micro-whip scorpions) collected in a 12 month period in the soil (0-
7 cm depth) of a secondary upland forest (120.1 ± 50.8 ind./mVmonth) and of a primary upland forest
(29.4 ± 20.2 ind./mVmonth) near Manaus all belong to the species Eukoeneniajanetscheki Conde 1993.
About 75% of all specimens inhabited the mineral subsoil (3.5-7 cm depth) where monthly catches were
negatively correlated with temperature and moisture content of the soil. Females were almost twice as
abundant as males. The lack of a distinct reproductive period and the presence ofjuveniles and adults
(both sexes) throughout the year indicate a plurivoltine mode of life. No specimens were caught on or
above the soil surface. Abundances ofE.janetscheki are compared withthoseofthe Schizomida(tartarids)
and Thelyphonida (vinegaroons) from the same study sites. E.janetscheki also represented the palpigrads
obtained from the soil of three other upland forest types in Central Amazonia (0-14 cm depth) and
accounted for 0.1-0.3% ofthe total arthropod fauna.
Terrestrial arthropods of Central Amazoni- Results presented are compared with abun-
an forests have been investigated for several dance data of E. janetscheki from three other
years (Adis & Schubart 1984; Adis 1997; upland forest types near Manaus, and which
Adis et al. 1997b,c) cooperatively betweenthe were sampled between 1985 and 1991.
National Institute for Amazonian Research
STUDY AREA AND METHODS
(INPA) at Manaus/Brazil and the Tropical
Ecology Working Group at the Max-Planck- Palpigrads were collected between 1981
Institute for Limnology in Plon/Germany and 1983 in the course of ecological studies
(Projeto INPA/Max-Planck). Data on abun- on Central Amazonian arthropods from two
dance and phenology of Palpigradi sampled previously investigated and fully described
during a 12 month period in 1982/83 in both forest types, all within 30 km of Manaus: (1)
a primary and a secondary upland forest are primary upland forest at Reserva Florestal A.
now available, as their time-consuming taxo- Ducke (= Reserva Ducke; 2°55^S, 59°59'W)
nomical evaluation has been completed (Con- on the Manaus-Itacoatiara highway (AM-010
de 1993, 1997). Our data represent the very at km 26; cf Penny & Arias 1982), (2) sec-
first contribution on the phenology of a trop- ondary upland forest at Rio Taruma Mirim
ical palpigrad species: E. janetscheki Conde (3°2'S, 60°17'W), a tributary of the Rio Ne-
1993. Voucher specimens have been deposited gro, where the vegetation was previously cut
at the Systematic Entomology Collections of but unbumed (Adis 1992). Both forests are
the Instituto Nacional de Pesquisas da Ama- subject to a rainy season (December-May: av-
zonia (INPA) in Manaus, Brazil, at the Mu- erage precipitation 1550 mm) and a “dry”
seum d’histoire naturelle in Geneve, Switzer- season (June-November: average precipitation
land and at The Field Museum, Chicago, 550 mm, but each month has some rain
USA. events; cf. Ribeiro & Adis 1984). The yellow
326
ADIS ET AL.—PALPIGRADI OF CENTRAL AMAZON 327
latosoil of the primary and secondary upland the soil and never caught on tree trunks or in
forests supported a 2-3 cm thick humus layer, the canopy. No specimens were captured in
interspersed with fine roots, and a thin surface ground photo-eclectors. In the primary upland
covering of leaf-litter. One ground photo- forest, palpigrads represented 0.2% and in the
eclector (emergence trap) and one arboreal secondary upland forest 0.6% of the total ar-
photo-eclector for trunk ascents (funnel trap) thropods extracted from soil samples within
were installed in both forests from December 12 months (Acari and Collembola omitted; cf.
1981 to December 1982 (Adis & Schubart Morais 1985; Rodrigues 1986). Their abun-
1984). Distribution of palpigrads in the soil dance in 0-7 cm soil depth was comparable
was studied between September 1982 andAu- to that of the Schizomida (Fig. 1). Most spec-
gust 1983 (Morais 1985; Rodrigues 1986). imens of E. janetscheki inhabited the mineral
Twelve soil samples were taken once a month subsoil (Fig. 2: 3.5-7 cm) and a few (22-
from each forest type at random along a tran- 26%) the organic layer (0-3.5 cm depth). An
sect with a split corer (= steel cylinder with average of 120.1 ± 50.8 ind./mVmonth was
lateral hinges; diameter 21 cm, length 33 cm) recorded in the secondary upland forest and
which was driven into the soil by a mallet. 29.4 ± 20.2 ind./mVmonth in the primary up-
The combined area of 12 samples represented land forest (0-7 cm depth). More than 50%
0.42 m^. Each sample of7 cm depth was then of the total catch in both forests was repre-
divided into two subsamples of 3.5 cm each. sented by adults (Fig. 2). Sex ratio (adult
Animals were extracted from subsamples fol- males to females) in the secondary forest was
lowing a modified method of Kempson (Adis 1:1.8 (81% ofthe total adults could be sexed).
1987). The monthly collection data of palpi- In the secondary upland forest, the monthly
grads from the two soil layers in relation to abundance of E. janetscheki in the mineral
changing conditions of precipitation, temper- subsoil (3.5-7 cm depth) was negatively cor-
ature and humidity of the air near the forest related with soil temperature, i.e., catch num-
floor as well as moisture content, temperature bers (in particular of subadults) decreased
and pH ofthe soil were statistically evaluated with increasing temperatures (total catch: r ^
with a linear correlation test (Cavalli-Sforza -0.7454, P < 0.01; adults: r - -0.5926, P
1972) using the original field data (cf. Morais < 0.05; subadults: r = -0.8614, F < 0.001;
1985; Rodrigues 1986) ofthe previous month. n — 12, respectively). In the primary upland
In addition, the presence of palpigrads in tree forest our data indicated a negative correlation
crowns of the primary upland forest was test- of adults with the soil moisture content in the
ed by fogging canopies with pyrethrum (with mineral subsoil (r — —0.5670, P < 0.\, n =
and without synergist) during the dry and 12). The total catches of specimens obtained
rainy seasons (July 1977, August 1991, Feb- during the dry season and the rainy season
ruary & August 1992, July 1994; Adis et al. were similar: 66% versus 34% in the primary
1984, 1997a). Palpigradi sampled were clas- upland forest and 61% versus 39% in the sec-
sified asjuveniles, subadults and adults (males ondary upland forest, respectively. However,
and females) according to Conde (1984a,b, there was no distinct reproductive period in
1993, 1997). the secondary forest (where E. janetscheki
was more abundant) becausejuveniles as well
RESULTS
as adults (both sexes) occurred throughout the
Palpigradi obtained from different upland year (Fig.3). These results indicate a plurivol-
forest types in the vicinity of Manaus were tine mode of life.
represented by only one species: Eukoenenia
DISCUSSION
janetscheki Conde 1993. The body length of
!
adult males (without flagellum) reached 1 Comparable data on the abundance and ver-
mm. tical distribution ofthe soil fauna in three dif-
A total of 146 specimens was collected in ferent upland forest types of Central Amazo-
the primary upland forest at Reserva Ducke nia were given by Adis and collaborators
and 599 specimens in the secondary upland (Adis et al. 1987a,b; 1989a,b; Ribeiro 1994).
forest at Rio Taruma Mirim. Out of these, Arthropods were collected to a soil depth of
92% could be identified to their developmen- 14 cm during rainy and dry seasons and ex-
tal stages. E. janetscheki was only found in tracted with the Kempson method as de-
328 THE JOURNAL OF ARACHNOLOGY
—
Palpigradi -- Schiiomida —“Thelyphonida
—
Figure 1. Distribution ofPalpigradi, SchizomidaandThelyphonidain the soil. Samples takenmonthly
at 0-7 cm depth between September 1982 and August 1983 in two upland forests near Manaus. (Total
catch = 100% in each forest type; n = total number of specimens). Total precipitation per month given
between sampling dates (= at the end of each month in the primary upland forest and in the middle of
each month in the secondary upland forest). The low rainfall observed in early 1983 was due to a strong
El Nino event (see Adis & Latif 1996).
ADIS ET AL.—PALPIGRADI OF CENTRAL AMAZON 329
Primarvuplandforest 90n Secondaryuplandforest
H-U2 N=541
11adultsimmatures 0adults immatures
(12,0 3 ”
%)/ \ £
-pw- /
1 1/ (63,A4%) W24,S6%)\ 1I0S •
o30-
J=juveniles S=subadults J=juveniles S=subadults
IS J A=adults A=adults
0-
0-3.5 3.5-7.0 0-3.5 3.5-7.0
SoilDepth(cm) SoilDepth(cm)
—
Figure 2.- Distribution of Eukeoneniajanetscheki in the soil according to soil depth, and percentage
ofall developmental stages in two upland forests near Manaus. (Total catch = 100% in each foresttype).
Samples taken monthly at 0=3.5 and 3.5=7 cm depths between September 1982 and August 1983. n =
total number of specimens.
scribed above. Between 75% and 92% of all 14 cm depth), the abundance of palpigrads
arthropods were found to inhabit the top 7 cm was lower during the dry season (33% of the
when Acari and Collembola were included in total catch; 9.6 ind./m^) but higher during the
the total catch numbers and 69%-"84% when rainy season (78% ofthe total catch; 33.7 ind./
they were omitted. Data on Palpigradi E. m^) when compared to the top 7 cm.
janetscheki; see Conde 1997) are now avail- A third study was conducted in 1988 in a
able. primary forest on white sand soil, about 45
One study was conducted in 1985/86 in a kmnorth ofManaus (Adis et al. 1989a,b). Pal-
secondary upland forest on yellow latosoil at pigrads represented 0.1-0.2% of the total ar-
the INPA campus in Manaus, where the veg- thropods when Acari and Collembola are in-
etation was previously cut but unbumed (Adis cluded (dry season: 57.703 ind./m^, rainy
et al. 1987a,b). Palpigrads represented 0.2- season: 74.255 ind./m^) and 0.5“0.9% when
0.3% of the total arthropods when Acari and they are omitted from the total catch numbers
Collembola are included (dry season: 50.448 (dry season: 14.119 ind./m^, rainy season:
ind./m^, rainy season: 63.850 ind./m^) and 15.023 ind./m^). The abundance ofpalpigrads
0.9% when they are omitted from the total in the mineral subsoil (7-14 cm depth) was
catch numbers (dry season: 11.934 ind./m^, higher during the dry season (86% ofthe total
rainy season: 17.886 ind./m^). In the mineral catch; 57.8 ind./m^) and during the rainy sea-
subsoil (7~14 cm depth), the abundance of son (89% of the total catch; 120.3 ind./m^)
palpigrads was higher during the dry season when compared to the top 7 cm.
(62% ofthe total catch; 62.6 ind./m^) but low- To which depth E.janetscheki occurs in the
er during the rainy season (44% of the total soil of the Central Amazonian upland forests
catch; 72.2 ind./m^) when compared to the top is unknown. First studies below 14 cm soil
7 cm. depth in a primary forest on yellow latosoil
Another study was made in 1990/91 in a and on white sand soil 45 km north ofManaus
secondary upland forest on yellow latosoil, showed that soil layers in 20-30 cm depth
about 50 km north ofManaus, where the veg- were dominatedby social insects, inparticular
etation was previously cut and burned (Ribei- Isoptera. Palpigradi were not found below 10
ro 1994). Palpigrads represented 0.1-0.2% of cm soil depth, probably due to handsorting of
the total arthropods when Acari and Collem- the soil samples (Harada & Bandeira
bola are included (dry season: 33.915 ind./m^, 1994a,b). In Costa Rica, palpigrads were
rainy season: 19.696 ind./m^) and 0.4-0.6% found during the rainy season in 15-20 cm
when they are omitted from the total catch soil depth with abundances of 350 ind./m^ in
numbers (dry season: 7.180 ind./m^, rainy sea- a forest and 75 ind./m^ in a coffee plantation.
son: 7.777 ind./m^). In the mineral subsoil (7- They represented 1.4% (800 ind./m^) and
—
330 THE JOURNAL OF ARACHNOLOGY
Primary upland forest
Secondary uplandforest
40 nSubadults 100 Subadults
(0
I20 - g 50
0)
ma. p Q.
1 (n
HM i a
SONDJFMAM. J, JA n - SONDJ FMAMJJA
40 Juveniles 100 Juveniles
w w
i20H E 50 H
o
0Q). a.
(/) Ui
EL JIL ELI FIMIAiMn1JnJI AI n1r11 1 '—p-nF— Mn pA—n pMi1J1 nJ\nA1
^S O N D J S O N D J
1982 > 1983 < 1982 > 1983 <
—
Figure 3. Temporal occurrence of developmental stages of Eukoeneniajanetscheki in the soil (N/m^
in 0-7 cm depth) in two upland forests types near Manaus. Monthly samples taken between September
1982 and August 1983.
0.3% (188 ind./m^) ofthe total arthropods col- supported by another study in the primary up-
lected to a depth of 20 cm, respectively (Ser- land forest at Reserva Ducke, in which no
&
afino Merino 1978). In Israel, Eukoenenia specimens were collected in 20 baited pitfall
mirabilis (Grassi & Calandruccio 1885) was traps and in three ormore groundphoto-eclec-
sampled to a soil depth of 15 cm in pine and tors during a sample period of 12 month (Pen-
oak forests. The 71 ind./m^ represented 0.07% ny & Arias 1982).
of the total microarthropods obtained (Broza Palpigrads are considered to be hygrophil-
et al. 1993). In Californian pine forests paL ous, photophobic, euedaphic inhabitants of
m
pigrads were detected to a depth of 1.18 soils or troglobites (Conde 1986, 1996; Janet-
&
(Price 1975). schek 1957; Mahnert Janetschek 1970;
The absence of E. janetscheki in samples Gruner 1993). E.janetscheki was not found in
from ground photo-eclectors in Central Ama- the soil of man-made pastures (0-14 cm
zonia also indicates that the species is not ac- depth) adjacent to upland forests in Central
tive on the soil surface. This conclusion is Amazonia. One reason might be the low hu-
ADIS ET AL.—PALPIGRADI OF CENTRAL AMAZON 331
midity andhigh temperature ofthe soil around the manuscript. Berit Hansen (MPI Plon) is
noon, particularly during the dry season (Adis thanked for making the drawings. Dr. Johann
& Franklin, unpubl. data). The abundance of Bauer, MPI for Biochemistry (Martinsried/
E. janetscheki in the mineral subsoil of the Germany) kindly provided the backsearch of
primary upland forest at Reserva Ducke and literature on Palpigradi, covering the last 30
of the secondary upland forest at Rio Taruma years.
Mirim was influenced by both abiotic factors. LITERATURE CITED
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ACKNOWLEDGEMENTS forest during the rainy season. Stud. Neotrop.
Fauna Environ., 22(4): 189-197.
This study was supported by a grant from Adis, J., J.W. de Morais & E.E de Ribeiro 1987b.
the Max-Planck-Society forthe second author. Verticaldistributionandabundanceofarthropods
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