l' II I I' 1 I: 1\

cology
David J. Curtis and Glauco Machado

arwstrncn arc a common component of terrestrial environ ments and . as we in-

ltml to show in this d lil plcr. a re often <m appropria te s ubject for ecologica l
siUd ics. They are dense and diwrsc in many habitats. arc easy to detect a nd cot-
lel'l. and in many count ries (especia lly in temperate regions) arc reasonably well
knownl<lXOnomkally. Al th ough n great amount of ecological information about har-
vestmen ex ists in the IOrm of quantitative samplings throughout the world . few at-
tempts ha\'c lx.'Cn made to understand their genera l patterns of species occurrence in
both time and space. t\ ft crsc<rch ing a wide ra nge of synecologica l literature. we have
;Jsscmblcd here most of the in formation on species occurrences in different habitats
and their to-ocrurrcnccs in communities in order to discuss the emergent trends.
In the fir st two sections or this chapter we consider spatiul and temporal patterns
of spcdcs di stribution s. stressin g the major environmental fa ctors related to h ar-
vestman abundance and richness. In the thi rd section we present basic data on h ar-
vest ma n communities. with special reference to species richness and relative abun-
dance. troph ic rch1tionships. and resource parti tion ing. Finally. we discuss the impact
of human activity on harvest man populations by providing some examples in which
they are ncg<~livc ly or positively affected and mentioning some enda ngered species.

SPATIAL PATTERNS

Distribution and habitat use

Harvestmen appear to be ubiquitous. being found in all continents except
Antarctica from the equator to high latitudes (8 1N and 56S; Cokendolpher & Lao-
franco. 1985). We can find them in a great variety of habitats: in soil. moss. leaf
liner. and grassy clumps: under rocks. stones. and debris: on vertical surfaces from
tree trunks to stone walls: and running over tall vegetation (Cioudsley-Thompson.

280
 Ecology 281

1958). The most diverse harvestman communities are reported for tropica l areas
(Figure 7.1A), especially humid forests. In fact. 18 of the 20 communities reported
to have 20 or more species occur in tropical areas. Perhaps the localities with the
richest commun ities in the world are found in the coastal Atlantic forest of Brazil.
where the harvestman fauna commonly exceeds 2 5 species in a given area (Pint o-
da-Roch a. 1999: Pinto-da-Rocha et al.. 2005). There is a clear decline in species
richness toward the poles, and accordin gly, in tempera te regions it rarely exceeds l l
species in a community (Figu re 7.1A). This pattern of distribution has been widely
reported fo r many taxonomic groups. such as insects. arachnids. severa l vertebrates.
and plants. and may be related to ecologica l and/or phylogenetic fac tors (rcl'icw in
Wiens & Donoghu e, 2004: see also Chapter 3).

40
Temperate regions
A
~
35 0 Tropica l regions
:5 30
] 25 Figure 7. 1. (A) Histogram showing the
distribution of local richness of harvestman
0 20 species in tropical and temperate communities
(B) Box-ptot (median. quanlles. and maximum-
1: 15
minimum values) of the local richness of
s~ 10 harvestman species in forested (n = 89 sites) and
z open habitats (n = 70 sites) around the world
{Mann-Whitney, Z(U) = 7.539:p < 0.0001).
Sources: Europe: Todd (1949);Wil!iams (1962):
Pearson and White {1%4);Curtis {1973, 1975.
1-3 4-6 7-9 10-12 13- 15 16 - 18 19-21 >22 1978a,b}; Davis and jones (1978): Chemini {1980,
Local richness 1981): LOser (1980); Bliss et al. (1981): Bliss
(1982): Adams ( 1984); Almquist ( 1984): Bliss and
Tietze (1984): Hippa et al.(1984); Rambla (1985)
60 B Schaefer (1986): Decleer and Segers (1989): B!ick
and Bliss (1991): Owen (1991);Aiderweireldt et
al. (1993): Gruber (1993): Freudenthaler (1994a.b.
1999): RU!itka et al. (199S); Komposch (1996.
1997a.b, 2000a): Docherty and Leather (1997);
Komposch and Gruber (1999): loch (1999):
lingerie (1999): Dennis et al. {2001): Hicks et al
(2003). North America: Edgar {1971): Carter and
Brown (1973): jennings et al. (1984): Corey and
Stout (1990): Cokendolpher et al. ( 1993):
Koponen (1994). South Americc1: Cokendolpher
and lanfranco (1985): Capocasale and Gudynas
{1993);Gnaspini (19%): Pinto-da-Rocha {1999)
Adis (2002): Bragagnolo ar1d Pinto-da-Rocha
(2003: unpublished data): Brescovit et al. (2004)
Forested Almeida-Neto et al. (2006): L. E. Acosta
{unpublished data): A. B. Bona Ido (unpuhh,hed
Ty pe of ha bitat data). japan:Tsun.rsaki {1999. 2003)
282 Ecology

:\lilwugh lwrwstlll(.:n appL'<Ir to be more diverse in mesic envi ronm en ts. they
<LI"l' not uncoJlllllOil in xcrk L'IWironnK'nts. even deserts /Hunt. 1991: Cokcndolphcr

d ;1L. 1lJlJ ~!.The sdernsonwtid Tmtll.wllinns 111amwm111S. ror instance. may be lo-
ndly wr v <Jbuml<1nt in desert arL'<LS of the Chihu11huan Desert in the southern USA
~ 1\liKKii Y d ilL. 1LJ1)2L 1-lmvevcr. if we analyze the harvestman richness from
dillerL'Ilt typ(.:S of 11ilhilats. il striking pallern emerges: median richness or har-
\'es\Jncn in l"orcs!cd h<~hilals is l.H times higher than in open habitats. includin g
gr:1sslands. rocky landsr<Jpcs. deserts. gardens. and bogs (Figure 7.1 B) . Open habi-
t<Jt s show mt~rkcd SL'<ISOJ\il] Vil l"i alions in abiotic factors. mainly tempera lure a nd hu-
midit y. whid1 probably rcstrkt !he oecu1-rencc of many h;irvcstman species (see
L'lwptL'r 1-1- ). Additionally. l"on..:stcd lwbil<lls are structurally more complex and prob-
ably prmide <I gre;Jler din:rsity or suit<Jbk habitals
Some harvcs\Jmn occur in nlounlain and alpine habitats. being recorded as
high as -1- .000 min tlw Hilll ClhJyas ([V1artcns. 1984. 1993a ) or in the p<lramos or
Colnmbiu (Hoewer. 1Yl-1-b ). but species richness seems to decrease with alti tude. Em-
pirkill datil on altitudinal dist r ibuti on l"or harvestmen arc scarce. but our analysis of
llilfvt'Stmun wrtil'<li dislribution (9()()- 3. 300m ) in the Eastern Alps from Komposch
and l;rubcr 11 Y99) revecded a monotonic decrease in spec ies richness along the ele-
vation a] gn1dicnt (H 1 = ().:S 1: F = 4fJ.O I: p < 0.001 ). Moreover. species wit h higher
mean dev<~ti o ns \ended to rover broader elevational range sizes. thus inflaling the
sperics number at lower cle\' Htional bands. Species at higher elevations probably
lww more tolerance to dwn gC's in cllmatic factors. such as temperalure and hu-
midity. bcc<mse !he climate is more \'arii1ble a\ mountaintops. This pattern of distri-
bulion is known as "RapoporLcrrect" (Blackburn & Gaston. 1996) and has been de-
scribed for bullerllits (Fleishman et al.. 1998) and a nts (Sanderset a!.. 2003). In the
future. studies in tropical and small clevationa l ranges would help the understanding
of how harvcstrmm richness dmngcs along gradients where the variability in climatic
factors is less accentuated than in tcmpemte regions.
Regarding spatial distribu lion nnd hnbitat occupancy of individual species. some
harvestman species. especially those of the suborder Eupnoi. can have a wide distri-
bution and can be found in a wide range of habitats (see also Chapter 3). Perhaps the
most widespread harvestman species is the phalangiid Milopus morio. a Palearctic
species that occurs in North Africa, Europe, Asia north of the Himalayas. and Canada
and presenls a broad habitat ()Ccupancy, ranging from woodlands to tundra and from
sea level to high- altitude montane locations (Slagsvold. 1976: Bliss & Arnold. 1983:
Tchemcris et al.. 1998: Tchemeris. 2000). As would be expected. this species is not
uniformly distributed over its environmental ra nges and usually consists of a number
of disjunct populations inhabiting distinct habitats. Therefore. some phenotypic
differences have been found among populations of M. morio, as described by Jennings
(1983). Similar cases of morphological variation were described early on for certain
species of l.eiolmmnn (Sclerosomatidae) along a latitudinal gradient in the USA
(Weed. 1893b). There is a gradual increase in body size and leg length in individuals
of L. vitll!tmn. L. velllricoswn, L. politum. and L. a/driclli from the north to at least as far
south as latitude 3 7N. Additionally. the proportionate increase in the leg length
southward seems to be R[eatel:" than that of the body, Ac,..~r\m.?6u..1-'Vf>edf~.R..Q_..'\h.),th'i'w
 Ecology 283

most important factors determining this increase in si1.e are probably the climatic
conditions that, with the decreased seasonality and snow period in the south. pre-
sumably result in a more abundant food supply and longer period of growth.
Contrary to the examples just cited. many harvestman species. including most
Cyphophthalmi and Laniatores. are much more limited in geograph ic distribution
and habitat use (see also Chapter 3). This is especially the case wi th cave dwellers. or
Lroglobites. which are found among all suborders of Opiliones (Rambla & juberthie.
1994). The genus Texclla (Phala ngodidae). for instance. occu rs in ca\'cs from the
southern USA. and most of its representatives are considered to be troglobitcs (U bick
& Briggs. 1992: Figure 7.2A). Trog/opl1iles are those organisms in which some popu-
lations complete their life cycles inside caves. while others do not. The troglophilic
"Dag uerreia" /11ermis (Gonylcptidae). for example. occurs in severa l caves in sout h-
eastern Brazil bu t was also recorded from cpigcan habitats far from caves tpinto-d;~
Rocha. 1996b: F'igure 7.2B). Trogloxtms ure those organisms that usc l.'mes only
part of their lives. such as certain species of the genu s Gym (Sderosonutt id:u:).
wh ich enter caves to hibernate during winter snow in eastern Europe (Novak ct al..
2004). and species of the subfamily Goniosomatinae (Gony leptidacl. whkh usc
caves for diurnal shelter and duri ng reproduction (r..lachado. 20112).
Cave-dwelling harvestmen may exhibit some ;~daptations to subttrrunea n
habitat that genera lly are more evident among the troglo bites (Figure 7.2,\J. On t of
these adaptations is the reduction or ewn t'omplcttloss of the eyes (l'.g .. luberthic.
1964). Among some cavernicolous specics the on1larium is present. bu t no eyes art
visible externally: among ot hers the eyes art preSi.'lll. but the retina docs not haw
the characteristic black color and seems to be nonfunction al. Other rommon suhtl'r-
rancan features arc the increased length of the legs. partial or tott1l abSl'11l'l' of tegu-
mentary pigmentation. and weak sderoti1.alion or the l' Utide ((;oodniglll & <:ood-
night . 1960). Little is known about the physiological tldjustnwnt s or tlwst
cavernicolous animals. but they are probably Jess tolcrun t to changing conditions i11
their environmen t than rclatc.!d species living outside the l'il\'eS
Morphological adapl<ltions probably related to h:~hitat use may ;dso he fmuul
among noncavernicolou:s species. Spctk:s that liw mostly o n the \'l'gctntion prtstn t
a greater n umber of l<lrsa l stgments tlwn tht'i r rdatiwsthutli\'l'(Ul tlw ground. Th is
is the case for some arboricolous gonylcptids bdonging tot he suhli11nilies l'm lllpy-
ginac. Progonylcptoidcllinac. and Sodrc;minae. which lw w X to 12 sq! llll'll\ S in the
tarsi Ill and IV (Figure 7.20. Their doses\ relati ves of the suhliunilk!> I krn;mdiiri-
inac tmd Conyleplinae spend most of their lives on till' ground cuul the numlll'r of
tarsa l segmen ts in legs Ill und IV is always less th:lll X (Figure /.lilt. I! i!' possible thai
a greater number of segments helps h<~rvcs tmtn cling onto vegl'l<ltion hy fnrmi11g
coils at the end of their legs ((; uffcy el al .. 1000: see also Clwptl'r ll
Fine-sca le habitat prdertnccs may con trihull' lo IIll' distrihulinn of speck'!' h~
twecn microbabituts. with. for txample. difl(:nnt spcdl's sp~.:ndinp, lllllrt linll' in
different layers of vcgct<1tion !Edga r. I YI.JOJ. Thus smcdl. shtlrl-ll'p,gld s1wcks. sul'11
as trog ulids and nemas t onwtid.~ (Figure 7.2 1 ~). liw ill tilL' Sllil or ll'til' litll'f. ll'hilc
larger spedcs. incl uding the oligolnphincs. ;m~ found nwinly in Ihe grnund-layer
vcgetntion. Lo n ger-legged species ort:ur prednmin:m lly h itdwr up em tlw brandws tlf
Figure 7.2. (A) The phalangodid Texella cokendolpheri. a strictly cavernicolous species that presents elongated
appendages and marked eye reduction (photo: j. Krejca). (B) The troglophilic "Daguerreian inermis (photo: R. Pinto-
da-Rocha). (C) The gonyleptid lporangaia pustulosa, an arboricolous species that inhabits moist forests in
southeastern Brazil and has from 14 to 19 tarsal segments in legs Ill and IV (photo: B.A. Buzatto). (D)The
gonyleptid Neosadocus maximus, a ground-dwelling species, also from southeastern Brazil (photo: G. Machado)
Although this species is a dose relative of/. pusrulosa, it has fewer tarsal segments in legs Ill and IV (4 to 5), which
correspond to the plesiomorphic state and are re Ia ted to a terricolous habit. (E) The short-legged nemastomatid
Dendrolasma parvula. a species that lives in the soil and leaf litter in Japanese forests (photo: N. Tsurasaki). (F) The
long-legged Mitopus morio, which is commonly found higher up on the branches of shrubs (photo: j . G. Warfel).
Scale bars= 10mm
 Ecology 285

shrubs (e.g., Mitopus moria: Figure 7.2F) or on tree trunks and up in the canopy (e.g..
Ltiobunum rotundum and Pha/angium opillo). It is noteworthy. however. that the dis-
tribution of different species relative to environmental strata may depend on their
foraging activities and may also vary with time of day (Todd, J 949 : see a lso Chap-
ters 8 and 14). Moreover. the relationship between leg length and habitat use seems
to apply only for some North American and European species. but not for most trop-
ical Lanlatores. Thus. a lt hou gh widely reported in the opilionologica llitcrature. th is
relationship should be viewed wit h ca re a nd deserves rigorous testin g. l<lkin g into
accoun t the phylogeny of the group
Some harvestman species may also occur in very specific mkrohabitats. as is the
case fo r the gonyleptid Riose{flllldo bimbtni. which lives inside the nests of the leaf-
cu tter a nt Acromymux /obicomis. Maury and Pilali ( 1996J iiuggest that <HI I nests
provide a welter microhabitat that allows th is harvestman to colonize arid zones in
Argentin a. Tank bromcliads arc also frequen tly occu pied by ha rvestmen (e.g ..
Gonz<i lez-Sponga. 198 7. l 992b), probably because these plant s accumu la te water
and thus provide a su itab le microenvironment that prevents dimatic stress. Indeed.
some ha rvestman species arc enti rely dependent on ta n k bromcliads for reproduc-
tion. such as the gonyleptid Bour!fiiYitr a//lionmta (~la c hado & Oliveira. 2002: see also
Chapt er ] 2). Some triaenonychids. such as Musdcoltr picltl and Al!!idia l'irldata. ap-
pear to be restricted to moss. and their green coloratio n probably funt't ions <IS cn m-
oufl agc (Cioudsley-Thompson. 1958). t\ nothcr interesti ng example is t ht BraY.ilicJn
cscad abiid Bamlimn1S lilloris. which is mainly found in ti1L' inttrtidnl zone crmong
debris (Soares. 1979). As our knowledge of th e biology of har vc:>tmcn inrrtases.
many more exa mples of microhabitfll spedlkity will pmbc1bly be d i:;covcnd.

Influe nce of physical factors

The influence of physica l fac tors on the spatial dist ribution of han't'SIIll('ll has
rarely lx.-cn studied. Todd ( 1949 ) stated th<Jttempcn ttun.. and h utnitlity ttrt rlw mt,st
important determinant s of hnrvtstlll<lll distribution <111d Jwhil<lt liSt'. lin like scor-
pions cmd many spiders. harvest men arc wry susnptihle to dehydration. and the
need for moist habitats is probably a signilka nt et:olo!!ical ftlctor that limits the oc-
currence of most species of the order in xcrir zoms (Set' Clwpttr [4). Young nymphs
arc pct rt lcularly Vlrlncr<Jb(e to desicca ti on and thu s spend most of thdr titllt' in le;rl
litter. under logs cmd rocks. a nd in ot her moist minohcthitat s. t\s lhty grow. indi \id-
u als ascend to the veget ation. as was rcl'ordctl for ScVl'ril l S]X'tits of the fcrmilk's Sck-
rosomatidac and Ph alangiidttc (Todd . I Y4Y: Edgar. I Y7 I).
The most comprehensive study of the inllucnce of envirormKn tctl pe~ranlL'ttrs. StKh
as relative hum idity and tcmpcraturt. on har\'t-':\t nwn distribution was done hy lilgcu
( 197 1), who reporiL-'<1 laboratory rL':"Ults for fou r spcrk's nf /.J.'io/m111rmmnsistent \\'ith
their preferred minolmbilat s in the licld (seL' Tabk 14.11. /,do/muwu l'il/11111111- l'ittd
as one of the most widespread spc"ies in the nurt ht;rsiL'nl l !St\. living in <1 widt \'ark't y
of habitats from dense wtXMJI;tnd to open hogs- had tlw highL'St ttmptn tun prd~r
cnce a nd the lowest rclttlivc humidity prdi:rence. surviwtllongL'St in dry <tir. ;u1d lost
286 Ecology

lhl k:~sl 1\'L'i~h! under d esin;~tion. All he olher extreme. the corresponding valuc.,-s for
/.. po/irw11. :1 spelics vulnemblt to low humidity. arc consisten t wit h its life in litter in
l'l'l'.v moist situ:11ion s. /.. nr/mr. :uwtlll'r species found to he vulnerable to low humidity.
ol'tcn Ji\'l'S ill ecotones with l)penings in the cunopy thai aliNv penetration of rain and
dew. bu l :1void:-; sunlight by inhabiting litter. The l'ourth species. 1.. nlrlric/ri. had the
ltlll'cst IL'Inpen llll rl' prdi:rcn cc but w:1s intermediate in terms of rci<Jtivc humidity prcf
lrcnn::. ll'hich is consi:o;ll'nt wit h it s h1vorcd habitat or moist. cool shade under dense
1\'lllJdl:llld l'i lllllPY1vhere tire :sh n1h l:1ycr is spc1rse.
'l'hL int lmncc 11f habit:1t st ructure nn harvestman abundance and distribution re
mains poorly explored. Wo rking with <1 harvestman community in woodlands of
southc;Jstcrn Engl:md. Ada111S ( l YX-H round !hal leaves of different sizes resulted in a
Iill er l:tyer of wntrasting characteristics of density and compressibi lity. which in turn
favored the occurrence of diffe rent harvestman species. Thus the s h ort~ l cgged Nc>mas-
lmrm /JiiWinthrwm INenwstonmlidae ) W<IS associalcd with the dense beech litter. whi le
the long-legged /.ndnius l'pilippiullfs (Phalangiidae) wus most frequent in chestnut !iller.
which has [;ugcr intcrlcal' !i]J<ILL'S. Moreover. on the basis of the general distribution of
spl'Cies in woodl:mds. i\d:1ms ( l9S4) suggested thatlmbilat structure could be an im-
port<l lll rae! or alTecting h:lrvcstnlilll distributions. Edgar (197 ] ) also showed that
dill'crent harvestm<ln species <Ire round in diflCrenl vegetal ion types in woodlands from
Mich iga n. liS;\ . The results. however. may be attributed lo differential preferences of
the SJX'cics to the mil'rodinmlic conditions generated by each type of vegetation rather
lhtll1 u :i[X:Cilic il:i:iOl' ialion between ht1rvcstmen nnd vegetation characteristics.
Corey and Stout 11 990 ) showed that the abundance of harvestmen in sand hills
in Florid <I (USi\J was not conclated with shrub density. ground cover. or plant litter.
In unother study Jennings ct al. I 19X4) showed !hat litter depth had little innuence
on eilhcr mean catches or ground harvestmen or the mean number of harvestman
species. Thcrel'ore. more s tud ies a llied with experimenta l manipulation of the
habitat structure arc necessary in order to better understand the causal effects of
habitat structure on harvestman distribution s.
Un like to scorpions (Polis. 1990). harvestman distributions do not seem to be
greatly influenced by cdaph ic facto rs. In the case of thcTrogulidae. however. there is
a clear pattern of occu rrence in places where the soi ls are derived from limestone
and are thus rich in ca lcium carbon ales. Since rcpresentalives of this family feed ex-
clusively on snails (see Chapter 8)-ani mals that require calcium carbonate from
the soi l lo build their shells-the distribution of trogulids may actually reflect the
dislribution of their prey (Hi llyard & Sankey. 1989).

TEMPORAL PATTERNS

Phenology is concerned with the evident activity of a species over the course of
the year and usually focuses on recognizing when adults are active and mobile. A
trawl through much of the available literature has provided data for Table 7.1.
which lists the phenological patterns for 71 harvestman species in 16 countries.
!~~~~:,.;,~n~~asonal patterns of activity for a range of species in different parts of the world, indicated by ~ Locality" (states, in the USA). and in different

Taxa JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC ,_,_.,
CYPHOPHTHALMI
Parogovia Sierra woodland Legg & Pabs
pabsgamoni Leone Garnon, 1989

EUPNQI
Amilenus aurantiacus Austria montane Komposch,
1997a
AStrobunus laevipes Austria open Komposch,
1996
A taevipes Austria woodland Freuderlthaler,
1994b
Caddo agilis XX Japan woodland Tsurusakl.2003
Centetostoma Spain woodland Rambla, 1985
bacilliferum
Dalquestia formosa xX XX xX Texas grassland Cokendolpher &
Slssom.1000
Euzaleptus minutus XX XX xX XX Himalayas montane Martens. 1993a
Gagrella varians XX XX XX XX Himalayas montane Martens. 1993a
GIOblpeS sp. XX XX xx XX New Mexico montane/ COkendolpher
wood etal., 1993
Gyas annulatus Austria montane Komposch.
1997a
G. annulatus czech Rep. woodland Borek, 1958
Harmanda kancn Himalayas montane Martens. 1993a
XX xX XX XX xX Himalayas montane Martens. 1993a
H mgrotmeata XX Himalayas Martens. 1993a
H1ma/phalang1um XX Himalayas montane Martens. 199Ja
do/poense
H nepalense XX XX XX Himalayas Martens. 1993a
H. pa/pale xX XX Himalayas montane Martens.1993a
H. suzukii XX Himalayas montane Martens,1993a
Homalenows Spain grassland Rambla, 1985
qvaclfldentaws
H qvacJn(lentalus XX xX xX XX XX Spain woodland Rambla, 1985
fl'mu/mlt'I/J
Table 7 .1 continued

Taxa JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Locality Enviro nmen t Source

Lacinius ephippiatus scotland woodland CurtiS. 1978a

L ephippiatus Austria opeo Komposch.

1996
L. ephippiatus Austna woodland Freudenthaler.
1994a,b

L. ephippiatus XX England gra ss/wood Pl1illipson.1959

L. ephippiatus xX Germany woodland Schaefer. 1986

L. ephippiatus wales Pearson&

Wllite.1964

L hOrridus Germany grassland Bachmann &

Schaefer.
1983b
L. horridus Austria woodland Freudenthaler.
1994b
Leiobunum XX XX Spain woodland Rambl a, 1985
blackwalli
L. blackwalli Germany woodland Bliss eta/., 1981
L. blackwalli Austria woodland Bliss, 1982
Lglobosum XX Japan woodland Tsurusaki, 1986
L.japonicum XX Japan woodland rsurusaki, 2003
L manubriatum Japan woodland r surusaki, 1986
L. rotundum XX XX XX XX Spain woodland Rambla. 1985
L. rotundum Germany woodland Bliss eta/.,1981
L. rotundum Germany grassland Bachmann&
Schaefer.
1983b
L rotundum Austria woodland Bliss, 1982;
Freudenthaler.
1994b
L rotundum England grass/wood Phillipson, 1959
L rotundum Wales montane Pearson &
White. 1964
L rupestre AUStria woodland Freudenthaler.
1994a
L. townsendi XX XX XX XX XX XX New montane Cokendolpher
Mexico etal., 1993
Leiobunum sp Austria montane Komposch.
1997a
Lophopilio Austria 'ocky Freudenthaler,
pa/pinalis 1999
L. pa/pinalis Germany grassland Bachmann &
schaefer,
1983h
L pa/pinalis Germany woodland BaChmann &
"" "" SChaefer,
1983b;
SChaefer.
1986
L. pa/pinalis scotland woodland cunis.1978a
L. palpinalis England grasS/Wood Phillipson, 19S9
L.palpinalis Wales montane Pearson&
White, 1964
Megabunus Scotland woodland Curtis, 1978a
diadema
M. diadema Wales montane Goodier, 1970
M. diadema ,x England woodland Phillipson, 1959
Metopilio sp "" .X"" New montane Cokendolpher
Mexico etal.,1993
Mitopus glacialis Austria montane Komposch,
1997a
M. morio SCotland woodland curtis, 1978a
M. mono Germany woodland Bachmann&
"" "" SChaefer,
1983b
M. mono Austria woodland Freudenthaler,
1994b
M. morio Austria montane Komposch,
"" 1997a
M morio wales Goodier, 1970
M mono England grass/wood Phillipson. 1959
M mono Norway woodland Solem &
Hauge, 1973
M mono Wales montane Pearson &
White,1964
lt'lmlimttd)
Table 7 1 Continued
,.,.. JAN FEB MAR APR MAY JUN JUL A UG SEP OCT NOV DEC Locality Environment Source

Mitostoma scotland woodland CurtiS. 1978a

chrysomelas
M. chrysomel8s Austria woodland Freudenth81er.
19948
M.chrysomelas Austria Komposch.
1997a
M. chrysome/85 Wales Goodier, 1970
M. chrysome/8s XX XX XX X Netherlands grassland Meijer, 1972.
1984
M. chrysomelas Wales Pearson &
White, 1964
Nelima genufusca XX woodland Tsurusaki,2003
N. genufusc8 XX XX woodland Tsurusaki,2003
N. semproni Austria open Komposch.
1996
N.semproni Austria woodland Freudenth81er.
19948
N. suzukii XX ><X XX Japan woodland Tsurusaki. 2003
N. suzukii Japan woodland Tsurusaki,2003
Odie/Ius aspersus Japan woodland Tsurusaki, 2003
0. aspersus XX xx Japan woodland Tsurusaki, 2003
o. troguloides xX Spain grassland Rambla, 1985
o. troguloides Spain woodland Rambla, 1985
0/igo/ophus h8nseni XX Spain woodland Rambla, 1985
0. hanseni XX England grass/wood Phillipson, 1959
o. hanseni scotland woodland Curtis. 19783
o. tridens SCotland woodland Curtis.1978a
o. trldens Austria open Komposch,
1996
O. tridens Austria rocky Freudenthaler,
1999
o. tridens XX Germany woodland Bachmann &
Schaefer.
1983b: Bliss
etal.. 1981
o. tridens Germany grassland Bachmann&
Schaefer,
1983b
o. tridens Austria woodland Bliss, 1982;
Freudenthaler,
1994a,b
0. tridens Austria montane Komposch,
1997a
0. tridens Wales montane Pearson &
White, 1964
0. tridens XX England grass/wood Phillipson, 1959
Opi/io dinaricus Austria montane Komposch,
1997a
0. ravennae Germany woodland Bachmann&
Schaeffer,
1983b
0. ravennae Austria woodland Bliss, 1982
O.saxati/is scotland woodland Curtis, 1978a
Paranemastoma Austria montane Komposch,
IJicuspidatum 1997a
P. quadripunctatum Austria rocky Freudenthaler,
1999
P. quadripunctatum Austria woodland Freudenthaler,
1994a,b
P. quadripunctatum Austria Komposch,
1997a
Paraumbogrella xX XX XX XX Japan woodland Tsurusaki,2003
pumilio
Paroligo/optws scotland woodland Curtis, 1978a
agrestis
P.agrestis XX England grass/wood Phillipson, 1959
P.agrestis wales Pearson &
White, 1964
P.meadii scotland woodland Curtis, 1978a
Pha/a ngrum opilio Spain grassland Rambla, 1985
Pooilio XX XX Spain woodland Rambla, 1985
Pop!IIO Austria open Komposch,
1996

(Um lurwr/)
Table 7.1 continued

Talca JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Localit y Envi ronment source

P.opilio Austria woodland Freudenthaler.

1994b
P.opilio Austria montane Komposch,
1997a
Platybunus xx Austria Komposch.
OUCephalus 1997a
P. bucephalus XX Germany woodland Bachmann &
Schaefer.
1983b;
Schaefer,
1986
Pokhara XX XX xX Himalayas Martens.
occidentalis 1993a
Protolophus XX XX xx New Cokendolpher
slngularis Mexico etal., 1993
Psathyropus XX xx Japan woodland Tsurusaki, 2003
tenuipes
Rilaena triangularis scotland wOOdland Curtis. 1978a
R. triangularis Germany woodland Blissetal., 1981
R. triangularis Germany grassland Bachmann&
Schaefer.
1983b;
Schaefer,
1986
R. triangularis Austria woodland Bliss. 1982;
Freuden
thaler, 1994a
R. triangularis Austria woodland Freudenthaler.
1994a
R. triangularis Austria montane Komposch,
1997a
R. triangularis XX England grasstwoOO Phillipson, 1959
R. triangularis Norway woodland SOlem &
Hauge, 1973
R. triangularis Wales montane Pearson &
White.
1964
Sclerobunus New mootane Cokendolpher
robust us Mexico etal.,1993
Trachyrhinus New montane Cokendolpher
marmoratus Mexico etal., 1993
T. marmoratus Texas desert MacKayet al.,
1992
Xerogrella XX xx Himalayas montane Martens, 1993a
dolpensis
Zaleptiolus XX XX XX XX XX xX XX XX XX XX Himalayas montane Martens, 1993a
implicatus

DYSPNOI
Anelasmocephalus XX XX XX Spain woodland Rambla,1985
cambridgei
A. cambridgei Germany grassland Bachmann &
Schaefer.
1983b
A cambridgei Germany woodland Bachmann&
Schaefer,
1983b
Dicrano/asma XX XX l!lS XX XX XX Austria woodland Gruber, 1993
scabrum
Dicranopa/pus Austria Komposch,
gasteinensis 1997a
lschyropsalis Austria rocky Freudenthaler,
hellwigii 1999
1. hellwigii Austria woodland Freudenthaler,
1994b
I. kollari Austria Komposch, 1996
scotland WOOdland Curtis,1978a
b1macu/atum
N. bimacu/atum Wales montane Goodier, 1970
N. bimaculatum XX xX xX XX England grass/WOOd Phillipson. 1959
N. bimaculatum wales montane Pearson &
White. 1964
N /ugubre Austria rocky Freudenthaler,
1999
(Cimliuur r/)
 -
Table 7 .1 Continued

JAN FEB MAR APR MAY JUN JUL AUG SEP OCT NOV DEC Locality Environment Source

I--; /ugubre Germany gmsslrmct Bachmann&

Sclmclcr.
1983b
N./ugubre Austna woodland Blrss. 1982.
FreudcnU,aler.
199llb
N. lugubre XX XX XX XX Nether grassland MeiJer. 1972.
lands 1984
N. quadripunctatum czech woodland Borek, 1958
Re11
N. triste Aus tria rocky Freudenthaler.
1999
N. triste Austria woodland Freudenthaler.
1994a, b
N.triste Austria montane Komposch,
1997a
Sabacon imamurai Japan woodland Tsurusaki,2003
s.makinoi XX XX XX XX xX XX Japan woodland Tsurusaki, 2003
Trogulus Austria rocky Freudenthaler,
nepaeformis 1999
T. nepaeformis Austria montane Komposch,
1997a
T. nepaeformis Germany grassland Bachmann &
Schaefer,
1983b
T. nepaeformis Germany woodland Bachmann&
Schaefer,
1983b
 T. tricarinarus Austria rocky Freudenthaler,
1999
T. tricarinatus Austria woodland Bliss,1982
T. tricarinatus Austria montane Komposch,
1997a
T. tricarinatus Germany woodland Bliss et al., 1981

LANIATORES
AcanthOpaehylus uruguay grassland capocasale &
aculeatus Bruno-nezza,
1964
ACU!ISomB XX XX xX xX XX XX XX XX xX xX Brazil Machado &
tongipes Oliveira, 1998
Auranus parvus XX XX xX xX xX xX Brazil woodland Frlebe & Adis,
1983
Dlscocyrtus sp. 1 Brazil woodland Mestre& Pinto-
da-Rocha, 2004
Eucynortula lara XX XX xX xX xX Brazil woodland Frlebe & Adis,
1983
Goniowma xX xX xX xX XX XX XX xx XX Brazil Gnaspinl. 1995,
spe/aeum 1996
MISChonyx xX xX XX xX xX xX xX xX xX Brazil woodland Mestre & Pinto-
cuspidatus da-Rocha, 2004
Pachy/Oidellus XX XX XX XX xX XX XX Argentina rocky Acostaet al.,
gol1ath 1995
Tncommatlnae Brazil woodland Mestre & Pinto-da-
Rocha, 2004

Recoroed ac tiVuy 1n each month IS inclicated 1n uppercase for adults and lOwercase for nymphs. High leYels of acttvlty are marked with boldface. and strons peakS are underlined. The
symbol -- lndiC3tes no Obsefvauons Not all authors recorded nymphs.
~96 Ecology

By t'onsidl'ring in how nmny lllonths mlult s were recorded. a full range of season-
a lilies is appart'll!. fro111 L'Xln:nll'ly sleiH K"hrone (in only 1 month ) to completely
o.uryl'l1ronc tin ;d l 12 111 ont h:-:!. Very kw species co uld be regarded as n onseasonal
ll'urychro iH'l: olll' Cyphophthal111i. sis Eupnoi. one IJyspnoi. and seven Laniatores
tTahk 7.11. llmwwr. it is important to stress th at the number of eurychron e
sptr its prcstntetl in Table 7. I is clee~ rl y underesti mat ed. sin ce th e great majority of
!Ill' NLotropind l.anh1tort'S !1\tlest re & Pint o-d;J-Rocha . 2 004; G. Mac hado, pcrs.
nbs.). as 1\'dl as most l'yp llophthalmi ((;. Giribct. pcrs. comm .) arc present
t h rtlughuut till.' yeclr. but tlwre Cl i"C fe w st udil's on t he phenology or these groups.
It has lung been recogni~ld that differe nt ha n estnmn species lmve different
typts of Iill cyriL'S. Tod d I I ~4':>), for in stance. recognized three main life-history pat -
tern s for Bril ish harvestmen: (II) spec ies ar tiw till year with overlapping generations
and \'aryi ng J>LTinds of <~dull presence. ({J) ;umual spcdcs that overwinter as ny mphs
from egg:-; tha t hatrh in till' a utu mn. or lc) annua l species that ha tch in sprin g and
dk' ill.'ftlfL' winttr. th rough whkh they survive as eggs. The duration of developmen t
r:1n be a nnu;1l. bicnnhd. or perennial (i'vlart cn:-;. 19 7Sb). and the peri od of matin g
Ige nerally cquiv<ilt..n t to the L-crorJing o f active adults in collections) ca n be eu-
rych rnne or stt..nur hrunt... If it is stenrn:h rone, peak activity may be in the spring. or
in :-;u mnwr to autumn. or in t he wint er. In a more recent s tudy Tsurusaki (2003)
provided <~not her dassilkatitm of lutrvest man life cycles based on the hibernating
s t<~ ge t)!' 3tl speries from jupan . According to him. there arc spec ies 1/Hll hibernate as
ttll eggs 112 spcl'ies), (/J ) ju \'cn ilcs ( 3 species/. {e) adults (3 species). and (d) eggs and
ju\'Clli les (2 SIX't..'iCS). The s tudy also s tressed the relation sh ip between life cycles and
phylogeny. showing that . wi th few exceptions. the stage or hibernation is rather uni-
form within a fcunily or genu s.
There ;:m~ also variations in life cycles amon g harvestmen living in tropical a nd
ncar-subtropica l part s or the world. In Sierra Leone all instars of the Cyphophthalmi
Hml!JIJI'itr JHIIlS!JIIriJOIIi ( eogoveidae) occur th roughout the year. but show three
ptaks: one in jmumry [early dry season). one in April- june (early wet season ). and
one in September-October (i<JlC wet season). Lcgg and Pabs-Garnon (19 89) related
these peaks to changes in soil cmd litter decomposition associated with the activities
of decomposer orga nisms. In the wet cen tral Amazon ian fores t of Brazil , two species
of l.<miatorcs. ilurtmus JH!rl' l/.'1 (Stygnidac ) and Eucyuorll/111 IIIIa (Cosmetidae), have
life cycles adjll sted to the anmw. l llooding regime (Friebe & Adis. 1983). Reproduc-
tion and early developmen t occur while the forest is flood ed (probably on tree
trunks). and when the water recedes. development to adulthood proceeds on the
forest floo r bdorc the harvestmen retreat once more up into the trees. Acosta et al.
{199 5) provide an additional example with the gonyleptid Pacl!yloidellus golimll.
which occurs in the Pampa de Achala (ca. 2.000 m altitude) in Argentina. At this al-
titude there ure marked seasonal variations. especially in temperature, with a frosty
period from March to November: hence the activity of P. golial/1 is restricted to the
wu rmer part of the year (October-April).
The net effect of the differe nt ways in which species are influenced by environ-
 Ecology 297

mental conditions is that their activities are spread over different times of the year.
With regard to adult activity, nine out of ten species studied by Phil lipson ( 19 59)
were stenochrone. overwin tering as eggs. The only eurychrone species. with adults
being present all through the year. was Ncmastoma /Jimacu /awm (Nemastomatidae).
which contrasts with Curtis's (1978a.b) account of N. /)imaculatwu in western Scot-
land. where the species was more stenochrone. It is in teresting to note that in the
Durham populations this species overwinters as both eggs and udults: overwintering
eggs are laid from August to early December and hatch from April onward. but over-
wintering adult females have developi ng eggs that arc laid from fl.lay to July before
these adu lts die by August.
Phenological patterns may also present marked variation along an altitudinal
gradient. as was demonstrated by Martens (1984. 1 993a) for some sclerosomatid
harvestmen in the Himalayas. Figure 7. 3 s hows annuallife-historydiagrnms for live
species for which there are clear differences in their phenology. AI the lowest alti-
tudes Gayre/lt1 varim1s adults are active all through the year and have overlapping
generations. Eggs arc produced in March and Apri l. and th en nymphs can be found
from May to September. At a slightly higher altitude a similar pal! ern emerges. but.
as shown by Harmmu/a iiiSfmcla. generations overlap in March instead of September.
and nymphs persist alongside the adults for most of the yea r. Then. ns we go higher
into more extreme conditions. the overlap of generations disappears. <llld the period
of adult activity shortens. In addition. during the harsher parts of the year popuhl-
tions persist as nymphs (Hamumda l!i!f!'Oiinema). nymphs overlappi ng with <Jdults
(1-filll(l/p/w/anyiWII/II'JW/t'I ISr). or eggs (HiiiU!/p/W/11/I!Jill/11 c/o/ptWIISC).
It is clear. therefore. that the phenology of a species m<lY vt~ry acrus:; its g.co-
graphic range and also among the habitats it exploi ts_ In order to investig<IIL' this
subject in more det<til. let us come b<H.'k to Table 7.1 and examine the 2h
stenochrone species that occur in at least two different habit<Jts <llld s holl' v; 1ri;1tions
in their life cycles. Their habitats were dassilied using lOur broad eu tcgories: ((/1 open
habitats. including grasslands and moors/ he<1ths: (/l) woodl< llld lwbitats. which in-
clude deciduous. coniferous. and mixed l'orests. including snub: ((I rocky are<1s and
deserts: and (d) montane habitat s. Then. to test for <IllY effects of the lwbitat on the
dun1tion of adult phase. for C<lch species the number of months iiS <Ill Hdult ll'iiS
compared through a p<lired t-test between popuh1tion s in ead1 type of lwbil<~l
(wh ich co ntrols for ;my phylogenetic effects). The main result was tlwt wuodl<uHI
populations spend more time as mlults than popul;llions 11f the same species living in
grasslands or montane habitats (p < 0.051. \Vc found no differl'lll'l' lwt1ncn wood -
land and rocky/desert populations. but sin re only eight spcLics lwd pupuliiliuns in
both of these IHibitat categories. this could be the result of sm:lll st~mpk size. Ckarly
in open habitats ;md at high ;lltitudes I here is m<1rked scason; ll v:ll'i<ltion inlmirnn-
mcnt;ll fa ctors. mainly tempcmture <HHI humidity: l'DtlSt'quen tly. in those h<~hit<Jts
the period of adult <1tlivit y is signifiLilntly shortened. <1/ld popul<Jtinns persist <I S q.~~s
or nymphs most of the ye<1r.
298 Ecology

D E~g:-; D Juwnik~ fTdi Adults Overlapping generations

1-/iiiiii/I'JifllriiiSiiiiiiiiO//JOL'IISI'

3 100- -1000 m J--r-r--f--'-------'-----1

Figure 7.3. Phenology of some

Himalayan harvestmen. Note the
shift fromeurychroneadultsat
lower altitudes-even with
overlapping generations (black) and
rapid development from eggs-up 150 -2700 m
to the stenochronicity of the high-
altitude species, which spend much
of the year as resting eggs. The
period spent as juveniles also varies
between the species. Based on
Martens(19B4}.

HARVESTMAN COMMUNITIES

Species richness and relative abundance

Ecological communit ies can be described and compared using simple param-
eters. such as species composi tion. species richness. and relative abundance of their
representatives. In this section we usL>d data from 88 sites in continental Europe and
Britain in order to make some tentative comparisons between harvestman commu-
nities using such parameters (see references in Figure 7.4). These data are derived
 Ecology 299

100
90 A
80
~ 70
~

g~
60
50

0
40
Q 30
20
10
0
10 12 14 16
Local rich ness
2 1.0
B
Figu re 7.4. (A) Relationship bet'M'en local
richness of ha.vestmen and dominance

~$
~ (expressed here as the percentage of the most
!c 0.8 common species in the community) in 54
European communities. {8) Box-plot (median,
~ quartiles, and maximum-minimum values) of
., 0
thedominance(expressedhereasthe
proportion of the most common species in the

i
~
community) in forested habitats {n = 31 sites)
0.4 and open habitats {n = 23 sites), considering
only the European communities (Mann-
g Whitney, Z(U)"' 2.292;p "' 0.0219). Sources
:t::: 0.2 ReferencesarethesameasinFigure 7.1,
[ e)(cluding Pearson and White (1964): Curtis

0 ~------0-pe-n-------,-----F-o-re-st_e_
d -----,
(1973): Davis and Jones {1978): Bliss et at
(1981): Adams (1984}: Schaefer (1986}: Oecleer
and Segers (1989): Komposch and Gruber
Type of habitat (1999): Zingerle (1999): Dennis et al. (2001)

solely fro m studies that investigated commun ities Hnd 1hu s cxdlllk' :-:i nglc-s pcl'ics <IC
counts o r fauni stic lists. The sa mpk wrnpriscs a rd;Ltivc.ly smtdl LliLLlll)l'r of sp,,:cil's
( 53 ) belo nging to the su bo rders Dys pnoi t 191 and Eupno i 1 ~ 4 I. In spill' of t ill' hl'l
erogcn eily in sample quality. th e dat<L sl'l provid es us wi th <Ln npporl un ily 1o lind
general pa ltern s wi tho ut the compliration of int c rnmtincnt ;d VtLriation iLL Spl'lic:-:
compositio n.
T h e h arvestma n richness in the Europecm cnmLmltlitics analyzld lwrl' ra ng_ld
from 2 to 19 species. As previou sly recorded for scorpitu1 l'OLlllllULlilics I l'tLiis. 19<J lll,
there is an inherent tmd urtilir.:i;LI bio1s for species-poor l"OLllLmLn ilil'S to hl' rh: tr<tl'tcr-
i7.t!d by one dom inant species and for species-r ich ro mmunilil'S to sh o\\' LL\lll't' l'l'l"ll
ness (Fig ure 7.41\) . Dom inan ce te nded to ht higher iLL com mun itit's of n txn h ahi t<Lb
comptLrcd w ith communities of l'orested h<LbiltLI S !Fi g ure /A HI. Thi s p<tl\l'rLL is pruh-
300 Ecology

ably rl'hl tld tut\\'11 faclors: tutl'ollllllu nities of U jX' Il hiibit a ts have fewer species than
l'tllll llHIIIililsuf fores ted h<lbi t<lts, ;~nd (/J) conHnunilicsof open habitats may be com-
posed of ttiiL' or <1 few w ry n:sistant species that h uvc physiological Hdaptation s to
lokr;Jit' tht stn:ssful rond ition s of th ese habit<1ts and numerically dominate the
i\ \nrc than hO'){, of the npcn-habil<lt communities were dominated by
l'Oil HilLIHil y.
one of tht Ji,Jiuwing sptdes (H ll helongin~ to the fa mily Phalangiidae): P/w/mtnium
opi/io I I 7.1 ":.1. 0/i!fo/op/ws ll'irlmst l 7. 1%). Odit-1/w; lnl!lllloitlts (S.6%). i\lliloJJIIS morio
1X.fl'!.. L and Hi/ui'/Julriiiii!/Uiud.~ I H.6%). For at least one of these dominant species. I~
opilio. thcrt' is u det<Jiled physiolog icul study show in g that the individual s a re qu ite
rl'sist<1nt to wutl'r loss (Cli ngenpeel & Edgar. I ~6h). which may explain why this
specils inhabi ts and est<.Jblislws dense popu lat ions in open habitat s throughout the
\mrld
Thcrt has hL't'n no previous l'f10rt by opilionologists to determine wh ich factors
arc as:-;or.:i<~ l l'd with numLrka l dominance. but some trends seem to emerge from the
dataset on Europea n commu11i tks. First. large size is not a requisit e for dominan ce.
as has hcl'n previously reported l'or scorpions iniH1biting sand habitat s (Polis. 1990)
L;~rge SJX'ciL'S. sur h as Orlil'l/wi .~piiiOSIIS and J;gtWIIIIS C0/1\'('.\'IIS. arc generally rare.
1\'hcreas some snlilll species. such as Ntmtt.~lOma lugu/m.' and 0. lritlens. may be
rummon. Second. S]X'cialist fc..'(,"<lers. includi ng trogulids and some representatives of
the genus l srll.wvpsalis. whkh feed excl usively 011 sna ils (see Chapter 8). are rarely
amo ng till' dom inunt species. Third. domi nance seems to be habita t dependent. since
thr maj orit y or the species {I 3 ou t or 20) domin ate either in open or forested habi-
tats. hul not both. The seven species that domlm1tc in both habitats-Loplwpilio
JNtlpillalis. M. moria. NtmttSIOII /11 /Jimtwullll um , N. 111!]11/m, 0. l ridens. Pamnemaswma
lJIImlriplllll'l/1/um. and R. lfillll!lll/aris-Hiso dominate 7 5% of a ll sa mpled commun i-
tics. Their most readily apparent common characters are that they have wide geo-
graphic distributions and simply occur in a great variety of habitats. from wood-
lands to cultivated lields. suggesting that these features may also influence
numerica l dominance. l'v lorcover. dominant species seem to be highly fecund (sec
Table 12 .2). bu t this trend is not very clear and deserves further attention.
The studies on Europe;m com munities a lso provide interesting information on
the spatial and temporal variation in the relative abu ndance of individual species.
Two extreme cases of spatial variation are 0. tritlens. which ranged from 0.2% in a
wood from Austria {G ru ber. 1993) to more than 96 .4% in a wood from Germany
(Bliss. 1982 ). a nd M. mario. which ra nged from O.J % in a garden from England
(Owen. I 99 \ ) to 8 2.3% in a Scots pine fores t from Scotl and (Docherty & Leather.
1997). Why the relative abundance of these species varies so much along their dis-
tribu tion range is an important question to understand the structure of the commu-
nities. The answer is probably related to local ecological conditions, including abiotic
factors such as temperature a nd humidity. as wel l as availability of resources such as
prey and oviposition sites. Nearly nothin g is known about the role of intra- and in-
terspecific competition on the composition and relative abundance of species in har-
vestma n communities {discussed later). However. 8 of the 10 most common har-
vestman species in our dataset show a negative correlation between their relative
 Ecology 301

. Odiellus spinosus ~ Opilio saxatilis 0 Leiobunum b/ackwalli 0 Other

100

,._
~

"lS
.;'! 60
=
:J
80
I Figure 7.5. Long-term changes in the
relative abundance of 10 harvestman
~ 40 species in an English garden. Only 3
species (Odie/Ius spinosus, Opilio
"> saKatilis, and Leiobunum biackwalli) that
~ were present throughout the 10-year
"il 20 period are shown (after Owen. 1991). The
~ remaining 7 species (0/igolophus tridens,
Paroligolophus agresris, Mit opus mario,
Pha/angium opi/io, Opi/io parietinus,
1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 Leiobunum rotundum, and Nelima
gothica) were rewrded sparsely during
Years the studyandarepresentedas"Other."

abundnnces and the species richncl:>S of the ('Ommunities: in which Ihey arc found
This result su ggests that in communities where the resources arc somehow limited.
the <1ddition of species promotes intcrspecilk competition. which afftcts the rd clliw
ablJndance of their rcpresenlalives. including the numerically domi11<1nt spcdes.
Regarding temporal variation. one study reports iibundilllt'c data from cx<Ktly the
same traps over a full de<:iide. Owen I 1991 J concisely listed long-t erm changes in <m
En glish garden where there were marked ch<1nges in species composi tion <Jild thdr
relative abu nda nce from year to year. Only three spccfts \\'crt prtStntthroughout the
10-ycar period: Odi1/lus spino:ms. Opilin SII.Wiilis. and l.l'iolmmnn Markil'll/li (Figurt
7.5). The remaining seven species (()liyoloplws ll'ilkllS. Hrmli!loloplw.~ l!!fre.~li.~ . .\1.
morio. l~ OJJilio. Opilio paril'linus. L mlllrulum. and 1\',/imll not/rinrl were recorded
sparsely during Ihe study. t1lways wilh low relative abumlw1ccs. J\1nrtovtr. tht rda-
tive Hbundan(e of the three most common spcdes prts:tnlcd grc;rt 1ariatiun
throughout the study. Otlil'lllls SJ1i r w.~us . li1r instame. 1\'ilS cdways tht dominc1111
species. but its relative abundance mnged from 47.7% to HLJ. 5';: . llt'irrly <I t ll'llrtl]d dif-
fertn cc. The relative <lbundtm(cor f.. /1/ir,klla//i also presented <Ill enormous l'il ri<J iinn.
nmging from 1.4% to 11.1 %. Owtn (I ~911 dues not provide <111 expl<ill<ltion for sudl
annual variation. but the resu lts or this study indit'<~lt' that both SJX't'ies mmposition
and relative abundance may vary considerably owr time. The h'mpural I'<Jri<Jtiun of
species abu ndances certainly deserves more at tention because it nwy hl'lp us under-
stand how different species restxmd lo environmtni<J I dist urhann:.

Trophic relationships and resource partition

The importance of predators in energy ;111d nutrilnt l')'dinv. is<~ li1 nrt inn or thl'
quantity of prey biomass Ihey c..: apture. whit'h in lllrll is il l'ullt'linn or lhl pnti:Jtors'
302 Ecology

1kn:-;ity. bionw:-;:-;, nKtabolism. ;111d l'JH.!rgy tn1 nsfer ellil'iency (Polis. 1993). There is
littk inl'ornwtitlll on tlu:se points for har vestllll'll . and the duta presen ted here come
lioJHtlll' most n nn prehensi\'1.: :-;tudy on the role of Opi lioncs as predators (Schaefer.
JlJXhl. The study wc1s nmdm:ted in a hecd1 woody ecosystem in Germany. where
l.oJ'Impilio pulpiuulis. l..udui11S t'/JI!i/IJiilllWi. MilO/IllS morio. and P/aty/J/1111/S /maplw/us
ll'l'l't' the domim1nt species. Soil <Jnd litter sa mples showed tha t the total harvestmen
tinm<t tn int Sptrk's) hc1d a nw;m annual dtnsity of 19 individuals/m 2 and a mean bio~
Ill <ISS !<IS dry wl'i~ ht l of 1 I mg/m 2 . TheSl' values were very low compured with other
prt(tcJtnry mar roarthropods in the leaf Ii ii er. such as spidtrs ( 1 3 5 mg/m 1 ). staphylinid
lxl'lks tlf1 mg/m:J. ;1nd t't nli]X'tks ( 3h6 mg/m 2 ) . E.<iti nmtes suggest a n annual co n ~
sumplion r< tll' by tht to\<11 harvestman f;nmu nmging from 0.2 to 1.0 kcn l/m 2 /year.
l'l ll'St' \'idLJt:-; rtprestnt tmly 11 sm;dllhJCtion of the a nnu al production of the l e<Jf~litt c r
arthropod l'Ollllllunit y and only a small portion of the tot il l pressure exerted by prcda~
tory arthropods. incl uding s piders. staphylinids. and centipedes (Schaefer. 1986).
CoJil]lilrtJble result s 1\'CJe round in ;mother type of habitat. a deciduous woodlnnd
r( ll'l'St in En gi<II HI !Todd. I ~ .tlJ ). Tht number or harvestman species was 1 7. the den ~
si ty in tht grou nd layer fi lll ged throu ghout the year from 0 to 50 ind ./m 1 . and the
biomass ra nged fro m 0 to 200 mg/ m 2 Despite the lack of informntion on the role of
har\'l'stmen ilS predators in Todd 's article, it is possible to estim ate the annua l
Hnwunt of prey Cilplll rcd using monthly hctrvcstman densities (see Table 6 in Todd.
llJ4-lJ ) and adult d;Jity food mnsumption (nea rly 0.001 g: sec Phillipson. 1960a.b.
19fl31. The conclusion is that all har\'estmcn living on one hectare of land would
ha\'e <Jl'Ombincd consumplion of 6 5 kg of pre)' per year. By comparison. a similar cs~
tim att was made by Foclix (1996) for spiders. and the result was 47,500 kg of prey
in Olll' hcl'tart per ycc1r. However. it is important to stress th at Foelix's number ls
dearly overestimated. sin rc it docs not consider seasonal variation in density (of 130
ind ./m 2 no less). nnd it asswnc..-s a da ily rate of prey consumption of O. l g/day- 100
times more than that cstim <J ted fo r <Ill adult harvestman. Obviously, we need rig-
orous estimates of the impilCI of arachnids as predators based on cmpiricnl fi eld
d;Jta. since at present. and except for a study of desert scorpions (Polis. 199 3). the
availnble information is lmsed o nly on hypothetical calcu lations
Whenever we st udy spcdes assemblages. we may consider distributions a nd co.-
ex istence mechanisms in at least three ways: (a) tempornl activity (phenology) seg~
rcgation. (/1) sma ll-scale sp atia l segregation. a nd (c) resource partition. Nonover l np~
ping phenological patterns may be one of the most important coexistence
mecha n isms. and temporal succession of ha rvestma n species (such as that shown in
Figure 7.6 for Scottish wood l <~ nd species) is frequently seen. Harvestmen also sha re
their hnbi tats with ot her cursorial hunters. and these seasonal patterns may extend
to other art hropod groups. Indeed. in Scotlnnd. wolf spiders (Lycosidae) tend to have
thei r peak actlvity in spring or early summer. to be replaced in later months by h ar~
vest men (Stinglhammer. 1987). This ca n even be extended to include n onarachnid
taxa. such as beet les. LOser ( 1980). for instance. showed temporal separation be-
tween nocturn al Carabidae and Opiliones. in which maximal activity occurs before
and after midnight. respectively.
 Ecology 303

Figure 7.6. Seasonal succession of

Scottishharvestmen,shownasthe
percentage of the annual total derived
from two years pitfall sampling: all
species tothesamescale.Thetotal
counts for each species are Mitopus
mario:: 246, Lacinius ephippiatus = 298,
Nemastomabimaculatum = 11,241,
0/igo/ophus hansen!:: 160,
Paroligolophus agrestis :: 443.
Oligolophus tridens = 1, 123. and
Lophopilio palpina/is: 1,339.A sequence
of seven species shows successively later
peaks in adult activity through the late
summer into early winter. Mitopus morio
shows a fairly widespread activity period
with three peaks, and both 0. hanseni
and 0. agrestis are bimodaL The
remaining species show single, sharp
peaksinabundance.Thistemporal
sequencing may contribute to the
coexistence of these species in their
woodland habitat

Coexistence mechanisms may <t lso O!X' ra te on a spatial basis. so that SIX't:ks are
segregated by different habita t preferences. t\dnms ( 19H4. I 9H 5J, ftlr inst;lncc. fcwnd
hi gh ly overlapping diets and no cvidcme l'or resource competition in his study ol
harvestman communities in d!.'(.'iduous woods in England. !he nmi n spt.'t"il'S of
which were Aw:lasniOt"'/)lwlus mm/Jrid_qci. Nc1 1111Sit111111 Mlllllnlfulrllll. l..opl111pili11 pu/Jiilurlis.
l.trdlliii.S epl!iwimus. i\11. 1/Wrio. l.tiolnumm 11/athm//i. ami L nuwulw11. I k condudcd
!hat habitat structure and density were the nitical ft~c l ors inlluencing tht slruclurt' of
the harvestman communities. This condusion. of cmtrsc. c;umot be gencntlizcd for
other sites throughout the world. ;md the mechanisms of nx:xistl'llt'C of S[X'l'il's in h<tr-
vestman communities remain an open licld for fu turl' rcsLarch.

HARVESTMAN CONSERVATION

Human impact on harvestmen

Harvcstm;m distributions may be inlhrcnced by both biotic ;md abiotic l'altnrs.
including climiltic conditions and habitat struct ure. Thcrd(u-t. any hunHtn ill'l i \ ' il~
that promoleschanges in na lu wl habitats may ncga livdy :1fl(:rt Stl1lll' spt.>eit:-o of tlw
304 Ecology

group. I Iu man im pacts may be very subtle. and even the aclion of wa lkin g over
ground \'l')-ll'lilliun can an"cct spider and harvestman communities (Curtis et a!..
I'JYI)I. lllhlT human activities, hO\wVer. have tremendous impact on the habitats
n11 whkh lwrw:;tmcn arc dcpcndcnt. Agricu ltu ral a nd infrastructure development
pnljl'l"l:;. for instatH"l'.l"i lll result in habitat fragmentation, which cnn endanger ha r-
\'l'St llll'll dirl'l!ly or indirectly. In this section we list some stlJdy cases tha t illustrate
lww IHI1111lll mtivitics can ncgil ti vcly or positively affect harvestman popull;lliOns.
1\ I<illY rqmrts conCl't"ll th e impact of l"c rtili:~.crs and pesticides on harvestman
popul<1tions. In I ill' Czech Rcpublit numbers of the harvestmcm Plmlangiwn opilio
<lroppcd slightly with the use of urea+ amm oni um nitrate fertilizer for winter wheat
nups (1\'hkh was a lso as.o,;ociatc<l wit h a dra matic decrease in invertebrates overall ).
and thl')' <lroppc(l signilieant ly wi th the appl ication of deltamcthrin pesticide on
spri ng barley 1il'lds ( Pckti r. 19Y 7a. bl. Addit iona lly. spider and harvestman quant ities
were ;dso found to -be sli gh tly affected by combined fertili:~.er/ in scc tidde applications
on win ter wheat (Pek;\ret al.. 19 Y71. ThedTcctsof inseetiddeson harvest men living
in a pple ordwrds were studi ed in th e northwestern USA (Epstein et al.. 2000). and
the authors round signilic;u1t dccre;1ses in populations of many arthropod groups.
inrluding. a lthough usu ally to a lesser extent. harvestmen. It has been suggested
Ihal the long-legged hanestmcn stand hi gh enough not to touch the pesticide
covered ground. thus reducing the impact of con tact pesticides (Haws. 1995). 1f this
is true. short-lcgg(.'d species should be more severely affected than the long-legged
sptdcs. but this prr.,>diction has not been experimentally testt.>d.
Forestry operatio ns obviously can a lt er the habitat conditions available to har-
vest men. lnlhe USA Jcnningset al. (1984) found signiflcant ly more individuals and
species (mainly l.l'iolnmwn ca/mr) in unt ut strips and dense spruce-lir woods than in
dcc1r-cut strips. Species diversity of spider and harvestman communities in peat bogs
varil!s signilka ntly with he terogeneity in vegetation structure (Curtis & Signal.
1980). which in turn can be strongly inOucnced by management practices such as
drainage and peat cutting. With drainage or simply the passage of time. peat-bog
vegeta tion nmy be replaced by woodland (many peat bogs are also planted with
coniferous tree species). and consequen tly the opiliofauna changes. Sometimes the
actua l spec ies of trees that arc planted can have an impact on harvestman popula-
tions: rougher bark on tree trunks provides more refuges for ha rvestmen and their
prey and e<m influence the diversity of spider and harvestman communities (Curtis
& Morton. 19 74).
Less obvious. perhaps. a rc pollutant effects on forest habitat. Bliss and Tietze
( 1984) described changes in species composition in harvestman communities in
German woodlands affected by air pollution. The authors suggested that these
cha nges could be caused by a lterations in microclimate conditions. which in turn
were associated with changes in the vegetation. Another study on how pollutants af-
fect harvestmen has been done In a n abandoned lead and zinc-smelting complex in
Arnoldstein. Austria. which was in operation for almost five centuries. Rabitsch
( 1995) analyzed metal accumulation in several arthropod groups. including har
vestmcn and spiders. and showed that lead, the main pollutant of the region, be-
 Ecology 305

came heavily accumulated in all arachnid species. especially in those sites nea r the
smelting complex. Additional ly. sex-specific accumulation occurred mainly in har-
vestmen . with males usually having to manage higher qua ntities of this type of
stress. Finally. no indica tion of changes In life cycle pa rameters was found. but a
negative correlation between meta l accumulation and body weight was found. It
seems obvious. therefore. that soil and vegetation contaminated with metals may
negatively affect predatory groups. such as spiders and harvestmen . wh ich accumu-
late great qua ntities of pollutan ts from the lower levels o f the trophic chain.
Fire a lso has drastic effects on harvestman popula tion s (e.g .. Curry ct <1 l.. 1985. in
Australia : Loch. l 999. In Germany). The most compreh ensive study of the effects of
fire on arachnids was conducted by Schaefer (1980 a.b). who recordL>d the recolon l7.n-
Uon of a burned pine forest by sever<tl arthropod groups in northern Germany. Re-
garding the arachnids. o nly a few harvestmen and spid ers survived the lire. but thl'
burned area was rapid ly recolonized by individua ls origin atin g from surrounding
areas. such as woodln nds and open habitats. Aft er about two years th e burned pine
forest was characterized by a num ber of SJX.'Cics and a po pu littlo n density that did not
differ from the values in unburned pine forests in the vicinity. HoweVl'r. S!X.'Cics com-
position was markedly different because the immi grating indiv l du<~l s belonged to two
different ecological groups: (a) species typically found in the pi ne forests. such us the
harvestman Paroligoto, Jws agrrslis (Phala ngiidae). and (!J) species from open habil ats
that established dense populations aft er the fire. such a s the spider PanloStl tunubris
(Lycosidae). Thus the arachnids in the burned area formed a typical commLmill' quill'
differen t from the one tha t inh abited the origi nal pi ne forest.
Harvestmen may also be negatively ;1ffected by domestic an imals. Frlc-range do-
mestic birds. such as chickens and geese. resulted in some rL'<luction in har\'cstman tll'-
tivity in an experimental apple orchard in Michigm1.l1St\ (Clark & Cage. llJ971. whkh
might be expla ined as a result of direct predat ion by chickens ;md/or htlbitat distu r-
bance by both bird species. In <mother study the number of catcht..-s of machnids was
compared in three tussocky grassland arc<ts in upla nd Scotland: wi th no livestnc:k
with sheep only, nnd with sheep a nd callle (Denn is et ;d .. 2001 ). For nwst SJX:dl'S of
spiders. harvestmen. und pseudoSlorpions. the abu nda nce was greater in till' Lm-
grazcd area . However, for the lmrvestma n i\ liiOJJ/IS morio. the mnst com mon aradmid
species. significantly more individu;1ls were found in the shorter Sll'tlrds or th e tll't'tl
manngecl with sheep only. According to the authors. th is t>tltt crn or eaptlll'l'S rnay lmw
been an artifact or its behavior of walking over thl' grass ca nopy cmd thu s ahow th1..
pitfa ll traps used for samplin g in the Hrca with no livestoc k tmd !Hiler \'l'gtt tllinn.
But not all human impacts arc detriment al to lm r\'est men. ,\ n l'XIremc txamplt
of a benign effect is the construe!ion of artilicial islands and l hd r su hsl''lllt'tltl'uln-
ni?.<tlion by arachnids. as described by Komposch t l ~~ hi. \ Vith in t1h11Ut t hrt'l' Yt'< lrs.
fi ve species o f Opilioncs (Piw lrm!fiwn opi/io. Oli!toloJ11111 .~ tridC'IIs, l.twiuiust'jlllippilrtlls.. Is
tro/nmus /aevifX'S. a nd Nr li ma srmproui). as well as 51'! spider sp,._cks. had t'Stahlbhtd
populations in these tlrtilicia l isla nds. There arc sevcrc1l S,l' llil nth ropic SJX'dt'S. <I mil ht
conslruction of human settlements and bui ldi ngs l'<l ll pnwitk hnmts fi1r J'Mlpula litltl:-
a nd communit ies or harvest men. For exa mple. Novak l't <tl. tltltll I n'J)Ilrl lll<lltll' tiH
306 Ecology

h::; Sloven ian sprl'ic:;. 42 ill'l' I'O: Jund in urban cen ters. particular exa mples being Ph!}-
lmryimn i'J'iliP. Nclinmscrnprm1i. ()J!i/io sa.mtilis. Leinhlll!lml rolwulum. Troyulus nepac-
{iwmis. OJ!i/in paricti1ws. ilml Nmwstomu hiilml/l/11111 spnrwm. The richness of the
Sloven ian urban opiliofaun<l may be due to so me extent to the relatively sma ll size
ur these to\\'ns and the proximity or prirm1ry lmbilats. Certain species of Cyphoph-
tlwlmi ran thriw in disturbed environments. In Bioko. Equatorial Guinea, the
lll'O~IJ\Il'id I'WW/Ot'ia .\imnoiclcs could be li,und in l<.1rge densilies in a cocoa plantation .
Other species il!'l' iibund;mt ill Sl'lllimanaged bot<~nical gardens in South Africa. Sin-
g<~pore. ;md Sri L<ulk<JtG. Ciribet. pcrs.comm.).
Some symm thropic speci es 111<1Y increase their rnnges largely in response to
human activities. m<linly urb-.miwtion and agriculture. Plwlnnyium opilio and Opilio
parictinus tll'l' plwlangiids clmracteristic of disturbed habitats a nd widely distributed
throughout the Holarctic region: today. because of human inlroduction. they arc
also comnwn in New i'..t.'<Jland (Hillyard & Sankey. 19S9). Other well-reported cases
<Ire Opilio cww~lrinii tllld 0. ru:idwi IPha lan giidac). two synanthropic species that
<liT! expanding their r<111gcs in Austria (Komposch. 1993). and Lciolnmum /imlm/111!!.
<I spedcs that lives on houses and other man-made st ructures and is expanding its
n111ge in (:ermany (Bliss. 1990). Ncotropical examples include the gonylcplids lJisco-
1'!/l'llls im'tllidus. V. oliwrioi. and Mise/JOB!/.\' n1spidntus. which are found in both nat-
unll and urbiln environments (Elpino-Carnpos et al.. 2001: Mestre & Pinto-da-
Hnrh<l. 2004: Pereira ct al.. 2004) . These three species have wide geographic
distributions in Bmzil. contra.:'i\ing with ot her closely related species. which are gen-
erally con lined to some mountains or natural forest patches.

Endangered sp ecies
1\ s we have shown. many species of Opiliones seem to be currently threatened by
human act ivi ties that promote habitat degradation. destruction. and fragmentation.
Harvestman species that have very restricted distributions can be particularly en-
dangered if human act ivit ies unfavorably alter their habitats. This is the case for
most troglobitic harvestmen, which may be driven to local or complete extinction if
the cavcrnicolous habitat is disturbed by pollution, development of nearby land. or
even noncon trollcd visitation to the caves. Alltrog\obitic taxa arc considered to be
threatened in Brazil. but the o nly troglobitic harvestmen on the "Red List of Threat-
ened Species" are the gonylcptids fnndwnocmn uai. Pncl!ylospeleus strinalii. and Giup-
pol!ia clwrmsi and the minuid Spacleo/eptes spne/eusa. Another example is the phalan-
godid i\llniormts rmufoi in the Canary Islands. which has been found in only one cave
and is included in the "National Catalogue of Threatened Species" of the Spanish
government (Oromf & Izquierdo. 1994). Similarly. the troglobitic phalangodids
Tlxe/la rokm rlo/pl!eri. T reddel/i. and T reyesi, which are restricted to certain caves in
Texas. are listed as endangered species in the USA. In some areas these Texella species
are threatened by degradation of their habitats. and in others by the invasion of the
fire ant Solenopsis invicta into the caves (Reddell & Cokendolpher, 2001).
Opiliones found in very restricted habitats around desert springs or mountain-
 Ecology 307

tops may also be endangered if the habitat is being cha nged by human activities.
This Is the case for the pbalangodids Calicina minor and several species of the genus
Microcina, generally found under serpentine rocks. part icularly in association with
serpentine grassland or woodland vegetation in centra l Ca lifornia. USA. Al l these
species are being considered for listing as endangered species but have yet received
no governmental protection (J. C. Cokcndolpher. pcrs. comm.). Other examples of
vulnerable species include two Argentine gonyleptids: Podryloit/e/lusfulviymnulmus.
which is found only o n the top of the highest peak of the Sierms Chicc1s chain in the
province of COrdoba, and Pacll!)loirleJ/us bore/Iii, which occ urs as morphologintlly dis-
tinct populations in rapidly disappearing rain-forest patches in the provinces of Salta
and Jujuy. Indeed. given the high ra te <II which humans are destroying these rain-
forest patches. it is possible that some geographic varian ts of/~ lwrd/ii will disappear
in the near future (L. E. Acosta. pers. comm.). Up to now. howe\'er. no harvestman
species has been recognized as endangered in Argentina, and therefore. Pjllll'i!/l"flllll-
lallls and I~ !Jort'l/ii receive no governmenta l protection.
Opiliones. like other secretive animals. can be in danger of extinction wit hout
anyone ever noticing. Our knowledge of i\frkan h<rrvcstmcn is dos~.: to zero (see
Chapter 1). and the same is true for han,estmen of tropic<~l Asia and most or the
Amazonia. The process of lwbitat destruction in these regions has increased signi1i-
cantly in recent dec<Jdes. and we arc certainly losing tnany species of plants and cmi-
mals (Sayer & Wh itmore. 1991 ). in some cuscs even before their ronnal descriptions
by taxonomists. This may be partin1lttrly frequent mnong invertebrates. tspcria lly
those groups that have historica lly altrttctcd the <~llcntion or few rcscardll'rs. such
as harvestmen. Only recently have certain species-a lmost excluSi\'cly tht cm't'
dwellers- been receiving some attention of resct1n.: hers and nmscrvtttilmists ctnd
thus have some kind of governmental prot crt ion. Givcnt hc p;tucity nf trolu~ka l in -
formatio n for the great majority of harvestman spcdts. it is almost impossible tP
provide any comment about their nmservation sl<llus. Tlll'rcl"rm.:. pn:sLrvation nl
habit ats. instead of pi!rticular species. may lKthc llliU"t' ctlcctiw mtans 11f prcJit't"tiiJil
of the diversity of Opilioncs around th e world

CONCLUDING REMARKS

In the cou rse of this chapter we lookLd at spHiial and temp11rt1l p;llterns in till' mcur-
rcnccs of harvestman species in both large and s mtlll sn des. Tlll're;dkr. 1\'l' disntssLd
how the activities of some species arc distributtd uwr the l't"asmJs ;md toudwd em
how this may be inllucnt"t'd by various cnvirontnct11<11 liu:tors. \\'e u!so s< cl\' hllt\'
communities may be described and compared in terms nf such p:l rat mttrs as S!Wdts
richness and spedcs abundance. While \\'l' rtcognizt' till' markedly dilltfl'lll species
composition of geographica lly scpa nthxlmmtmmitks. we lw l'l' L'vidL'Ill"L' for at let~st
two potcnti<~ l mechanisms that nmlribute to the sep:1ration ami thus tlw l"lll"Xis-
tcnce of different species of lwrvcstnwn and othtr t<~xa: tL'rnpo ral acti1ity p<clll'l"tls
and habit at preferences.
308 Ecology

II is importantto tmphasizc. however. that there arc more than 0.000 species of
h<tneslmcn wnr!dwidc. li:w or whic h lw ve been reatured in this chapter. The ma-
itlrily 11r pttblic<tli<UlS rcnml the distrilnttionand taxonomy of single species . with no
itHlir<tlion or the coexistence of spedes in delimited aretJS. Thus we suffer from a
Sl'<trdt y of infonmttion tltthc community level. 1\t the population level there arc no
studies that prmidc {lala on su rvivorship. mortality. or any kind of intraspccilic in-
teraction s that may inlluence the distribution or individuals in th e space. There is
llllll'h In le;trn itboul these fasc imtti ng asscmbhtgcs or species. and because they, as
arthropod prcd<~tors. prob<tbly inlluence th e populations or many ot her species, we
have <ttnplc reason to expa 11d our knowledge of hctrvestman ecology. An updated
perspeLtivl' on IHtrvestnwn ecology was presented here. and we hope that it sti mu-
lates young arctclmo!ogists 1o develop <1 lifelong interest in the subject.

ACKNOWLEDGMENTS

We cxprL'SS our tlwnks to a ll our colleagues who were kind enough to provide us
with information und data. This chapter was great ly improved lhrough comments
by 1\. ]. S;tntos. G. Ciribet. (;.Q. Romero. M. Almeida- Neto. l~ R. Guimarf1es Jr.. R
l'into-chJ-Rocha. H. Mucias-OrdOi'lcz. and three anonymous reviewers. D. Curtis ded-
icates this cluipler to the Jute Dr. Ron Pearson. who. as his Ph.D. supervisor years
;tgo. inspired him to set off on this long voyage of discovery. and to his wife Angela
for her constant support. G. Machado is supported by a research grant from Ft\PESP
(02/003X \-0).