C H AP TER 1

What Are Harvestmen?

Glauco Machado, Ricardo Pinto-da- Rocha, and Gonzalo Giribet

T
he name Opilionts. proposed by the Swedish zoologist Kart 1. Sundt'\'<111 t I X~ Jl.
derives from the Latin word opi/io. us!.!d by the Ron1<111 dralllil!iSI Plaut us t2'54-
l 84 BC) in h is colllL"<Iits a n d mt;ming "shct't>-mastcr"-ont or tilL' Vil f ious etl ll'-
gorics of Roman slaves. Virgil's Edoy11es. wriltt'n in ~~ Bl'. ;llso lll!.!ntion s the 1\'(Jfd
opi/io. but alluding to <I slwp hcrd. This assod;~tion is prob;1hly rd;,tcd to the ell'\'tltl'd
body posi tion tliTortkd by tht long kgs of ccrt:. in har\'l'stman st>ccks. thus rt'St'lll
bling ancient Eurotx;m s hepherds who used to wantkr ;~bout on sti lt s in order to
better count th eir !locks. The l'<lrlit'SI al lusion to the order Opilione:-; in modern li!lr
aturc probtlbly occurred in '1111 '111!'1111'1" af lu~~ns by ~ l oflltt 11 h Hl. in \\'hkh tlwy
were ca lled sllcjJI/nd spitll'r.'i. a n a nll,' ~ till used in Crt'tll Bril<~in IJOII'illh1ys. The uuthcu
explained that people ccdlcd them slwph erds hera LISt' tiH:y tlHIU ~h t 1h<~l 1he Ikids i11
wh ich they wert abuJHlanttonstitu lt'll good s h tt..'p p;1sturt'.
T h e most pup ulc1r common namts in Engl;md tHIII"ilti<IYS :t rt /wnc~/111!'11 or,,,,,..
t'c~l.~pitll'rS. probably btcmi St' stunc speci es <~rcqui t c ;dlundanl illthtlwn"t.q St'<ISII II
An o ther hypo tlw~is is th a t th e convulsil'l' llltJ\'t'lllt'tl\ matk hy thl'ir lq!S <1lkr they
lmvc become dttadll'd from the tlll'llt'r's body til cwHIHml ddi. nsin hehadnr in
some species of th e order) rtstmhlcs tlwt or till" lwrwsllll<nl's sryt lll". Tht llll'<lll illg
of manycommonnwnts in European countries is ""rtapcr""tsct T<~hlc I. J 1. <IIHithis
a l ~tl mt~ y he rd:J tell to the harwst ptritM.I 11r ttJ!IJc <lpptarclnn ,,f rtapin g. :1s ll'itll :1
~cy t h e. w hen thcy wa lk . Curiously. in Stllllt' provi11n"sof Sp<1in hantstllll'll <lrt' tcllkd
JU'tlms in allusion to Seli n\ Peter's l>ay IJIIIlt' 291. whkh falls ll"ithin lht h:wwst
st:ason and is <~lime when some spcdts are t'S ill'd<~lly :~hundant inthl' 1idd:-.. , \pp<~r
ent ly the common names used i11 th l' Ne lhlTiilllds <IIH! Snuth ,\l"rkn. ll'hkh 111\'illl
""hay wagon " (see Tabk 1. 11. also <Illude to tht h ar w ... t period.
In l'Ollll lrics w hcrt: th r.: opiliolin11W is domillilll'tl by lun g-lq.!.).\l'tl :-.pl'l'il'.'-. lhl'
cornmo n1w nws gt:rll' rillly make rt'!lrenl't'to this L"lliii'<ICll'ristit lllnrph,Jio~k;d lr<lil
(;ood examples arc tlmlt/,11 lwrrJh'!l~ usl'd in North t\mnila :md .\u str:ili<l. <11"111111\
2 Wl l.:ll Arc Hilrvestrnen?

Tab le 1.1 Populm munes attributed to Opiliones around the world and their respective meaning in English

Name (singular) country (language) Meaning

Afterspmne Germany (German) False spider

Agostero Spam (Spanrsh) Flock of sheep that graze on recently
harvested fields. or simply harvestman 1
Aralia piltona Mexico and Spain(Spanishl Long-legged spider
Aranllaallro Brazil (Portuguese) Garlic spider
Aran/Ja-barlarma Brazrl (Portuguese) Dancing spider
Aran/ra-bode Brazrl (Portuguese) Goat spider
ArartiJafedorerJta Brazil (Portuguese) Stink spider
Bodum Brazil (Portuguese) Bad smell
caner united States (English)
Clricltina or chinclrina Argentina (Spanish) Sti nk bug
Daddy longlegs United States and Austra lia
(English)
Faucheur or fauclreux France (French) Reaper
Fradefedorento Brazil (Portuguese) Stinky friar
Frare Spain (Catalan) Friar
Giramundo Brazil (Portuguese) Traveler
Grandfather-graybeard United States (English)
Harry-longlegs United States(English)
Great Britain (English)
Hooiwa SouthAfrica(Afrikaans) Hay wagon
Hooiwagen Netherlands(Dutch) Hay wagon
Jangnim kOhmP Korea (Korean) Blind spider
Kanker Germany (German) After the Latin word cancer. which
means ~crab"
Kaszasp6k Hungary (Hungarian) Reaper
Kosar Ukraine (Ukrainian) Reaper
Kosarz3 POiand(POiiSh) Reaper
Kosec Slovakia(Siovakian) Reaper
Langbein Norway (Norwegian) Long legs
Lockespindel sweden (Swedish) Curled-hair spider
Lukki Finland(Finnish) word used only for Opiliones, with
no other meaning
Mangzhu China (Chinese) Blind spider
Matija Slovenia (Siovenian) Long-legged person
Medelwr Wales (Welsh) Reaper
Mejere oenmark(Oanishl Reaper
 What Are Harvestmen? 3

Meaning

Mekuragutmr Japan (Japanese) Blind spider

Miljdeci or miJcded Turkey{T\JrklshJ Person who brings good news
Pauk kosac or just kosac Bosnia and Herzegovina, Croatia, Serbia and Harvestman spider or just harvestman
Montenegro, Macedonia, and Slovenia
(Slavic languages)
Pedro Spain(Spanishl In allusion to Saint Peter's Day, which
matches with the harvest period
Pendejo Costa Rica (Spanish) Fearful
Pinacates Mexico and Spain (Spanish) Name used lor a black beetle that lives in
damp and shadowed places
Putnik Serbia and Montenegro (Serbian) Traveler
Schneckenkanker Germany (German) Snail harvestman
sekatt czech Republic (Czech) Reaper
Senokosec Russia (Russian) Reaper
Shepherd spider Great Britain (English)
suhajulina Slovenia (SiovenianJ Thin, bony, and mostly tall person
Teiliwr wales (Welsh) Tailor
Vevkjerringer NOfWay (Norwegian) Old woman weaver
Weberknecht Austria and Germany (German) weaver's helper
zatomushi Japan (Japanese) Blind bug
Zimmermann Switzerland(German) Carpenter

Names directly derived from the name of the order. such as opihonidi (Italian), opili6n \Spanisll). or op1hiio (Portuguese). were not uiCiudec! in tile list
1. The first meaning was extracted from Rambla {1974). and the second is an alternative hypott1ess snce tile word agosrero comes lrom agosto (August).
which is a harvest month in Spain
2. ln some places the word kOhmi (spider) may also appear asgumi
3. This word clerilles from kosiarz. which means "reaper."
4. This name is no longer used in Japan because the word mekura has dscnmnatOfY cormotatons to bhnd people.

pa!OIIfiS. used in Mexico. and Sfflw jfl:iw ami 11111/ija. USl'd in Slnwni;t rr:.hle 1.11. In
tropical region s. especially in South Alllcrint. whl're I he opiliol';rww i .~ rn<rinl y rom-
posed of robu st and short-lcggl'd .-;pl'rit'S, the common llilllll'S ill"t' u .~ tt<rlly rchrtl'd tu
th e had Smell rciC<ISl'd by Ih e seen I gl;uu]s lm"illl'd llll I he <rlll l'l"inr lll<rr~illS ol" the
body. <t unique morphological l"c:~lltrc of OpilitliH'S. J n ,\r~tntin:~lhl'y u stthl' 1\'urd:-.
d1idfi1111 or d1i11d1i1111. w h ich ;1re crlso crpplkd lo sli nk bugs or thL' onkr I il'll'ropll'l"it
!Tttblc 1.1 ). In Brazil names s uc h as /IO!f/1111. lfl"lllllfll-/lollt. onfllllrl-/t't/ormla. unmlro-
(11/w. ;.md frtrtlt-fidorl'lllo arc all rdatl'd to t ill' slron g sour snwll Sl'l"l"dnl hy tlw ];trgt'
gonyleptids ('J'ablctt. JJ, by far the moSintlllllltJil l"amily inlhl'l'IIUil lry
Biologisl s kn ow l i tt le a1")(1UI the tmilr ( )pi l itllll'S. l'\'l'tl tl ttlll).:lt il utsli l tlll':o-. :rfllr
the Acari (miles and licks) ;md i \nuwal' (spidl'rsl. lhl' lhinl-l<tr~L' S I ~roup or <tf<~t "h
nids. I n many cou ntries the gcnenrl pub1ir 1aq.!,ely ignorL'S h<tl"\'l'S IItll'n <IS mll. 1\r-
h aps because most species h ave stcretiw horhit :-~. lil'l' ir1 d;~rnp :~rtd sh<td(\1\'l'd <tl"l'il:-.
arc dH r k colored. and arc al"livc m<tinly ill 11igltt. thl'i r txisttnn paSSl'S nlnt~1:-.l llrlrl<l-
4 Wlu t A1e H<nvt.>stmen?

tintl b~ hlllll<ltl :\ . which 111<1)' explain th ei r virtu;il ab:-;cnce in myt hology. folklore.
;111d llisttll',l'. It is ll'twth tHitinl! tlwt Rt1hcrl l luukt (I M1)). the inven tor of the rnicro
.~orx. nHntion .~ in his hook Alifn'!/rtltJIIia ;mold supe rstition or the county of Essex.
En glaml . i11 n1l\'i ng h;n tst mcn. ,\t'cording to the tcJie. killing harvestmen on pur-
pust wuultl hring b<1d luck lxct~USt' supposedly these mystind c.minulls would help
the f<~rnlt'fs Jmr\'L'st the tTops wilh the scy the !hey IWI'l' alleged In possess. On the
other h;11ul. lwrvcs\Jmn do not h<1\'L' surh good f:une in the United Slates and ;\us-
trali<L ;md thtn is <1 persisttttt urlmnlegtml!halthey arc extremely poisonous. al-
though thdrn u Juthpil ri S<I I'l' ltiO ti ny to inllkt wounds on humans. 1\ syou will learn
in this !l!lok . lwrwstlntn <ll't' beautiful an1t'hnids that pose no d;lllgcr to humans.
Therl' art Sl'\'c nd explana tinll S of tlw origin ol' thi s ll'gcnd. butt he most plau sibk is
till' wn ru si1111 regarding tht llilllll' " dt~<ld y longlegs... which is also used for spiders of
till' fanti ly l'h clldtlill' in tllt lSl' l'IIUiltrit'S. Sinn S(IJllt' phokids regul<~ rly prey on ot her
spiders- indud ing the rctlhack spidtr (/.Jtlrmki'IWi /msst/li ). whose venom can be
ratal !II hulll<lll S- it iS poSSib lt thatthis ractlltlS Origirl<l!ed !hC rumor thai they arc
the mo~t diillgtrous spiders in I he word. Huwever. bel'ausc of their tiny fangs . phol-
t'ids. like lt;lr \'estmtn. art t't ltnpletely harmless to humans.
ln ttnns of gtnenl l morphology. lwrl'estmtn arc typical arachnids. They hai'C
two bt~ s i t' body regions. a pru:joma. wh idl t'il rrics all tht <tppcndagcs. <md a limbless
upisllwsnrn;L which hil s the spirt1dcs t~nd !he gcnilal openi ng. olicn tovcred by em
optrculum (Set Ch;1ptcr lL The jumLion between both body regions is notcon-
stril'tcd. giving tlwmllll' i!ppcarancc of "wa islless" spiders. Bc<'l.IUSC of Ihe superli-
cial rtsemhlarll't' hl'lwetn har wstrntn and !heir most famous cousins of the order
,\r;Hleae. Ollt' of their popuhn designation s in German is Afli'I'Sj1illllt', tiHrt is. "false
spider... t\nothcr common Ctr man name. I1'1'/ll'l'klll'dll ("weaver's helper" ), is also an
allusion to harvestman morphology: the second pair of legs is clon gatcx l in most
species of tht order a nd functions like an antenna. w;o~vi ng in the air while the
tlnirnc.JI is 1\'alking. thu s rtsembling the arm movements of a weaver's helper disen-
tangling th e threads on tl loom. (A nother meaning is "tailor's apprentice" because
harvestmen ca nnot lll<tke webs.) It is int eresting to note that the usc of the word
1\'i'/J('/'(Weavcrl has led to the wrong ide;.rthat harvestmen . which have no spinning
organs. n111 pr<xlucc silk.
Harvestmen are among the oldest anrchnids. and the fossil record demonstrates
that the group has remained al most unchanged morphologically over a long period.
a phenomenon called stasis (sec Chapter 5). Unique characteristics of Opiliones.
sud1 as p<.tired tracheae. the penis. the ovipositor. and the openings of the scent
glands. arc already observed in fossils from the Devonian , proving that the group has
lived on l;rnd and that males tra nsferred gametes directly to fema les as early as 400
mi llion years ago. In fa ct. harvestmen are considered one of the most primitive forms
of arachnid , possibly closely related to scorpions. pseudoscorpions. and solifuges; to-
get her these four arachnid orders form a clade called Dromopoda. However. the
exact phylogenetic position of Opiliones within the dass Arachnida remains a con-
tent ious issue in systematics (see Chapter 3).
The first harvestman species to be described were Plwlangium opillo and Trogulus
 What Are Ha!Vestmen? 5

tricariflatl/s, named by Linnaeus in 1758. Since t hat time some 6.000 species have
been descTibed by more than 110 taxonomists. of whom ha lf deS<.-rlbcd more than
10 species. Nearly 200 harvestman species scattered around the world were de-
scribed between the 1870s and the first decade of the twentieth cen tury (Figure
1.1 a), when Eugene Simon. Tore! Ta mer! an Thcodor Thorell. \>\Iiili am Sorensen
(Figure 1.2a). and Nathan Banks. among others. introduced more than 700 new
species. An intense descriptive period occurred between t he 191 Os and the I 9 50s.
when the German author Ca rl Fried rich Rocwer (Figure I .2b) was most nctive. de-
scribing more than one-third of the species in the order Opiliones. He created a new
system of classification. nnd his book Ok H'''xrki ii'Cilfl' der Enk. published in 1913. is
o landmark. cont ai n ing the 2.000 harvestman Spt'1.:ies known until tha t time and
many descriptions of new taxa. Rocwer's age was also a time of other important pro-
lilic taxonomists. such as the Brazilian C:i ndido Firmino de Mello-Lci t;:i o (Figurt
I .1c) and the South African Reginald Frederick L<nvrencc (Figurt: 1.1d]. SonH: Yt'itrs
lat er. between 1940 <IIHI 1980. the i\merican l"O ll pk rvl<trie Louise ;md Clarclll'l'
Goodnight (Figure 1.1e). the Brm:ilian HCiia Elllr l'vlontciro Soares~ Figure 1.11"1. and
the Japa nese Scisho Suzuki (Figure 1.1g) were also wry pnxluctiw. dlsaibing in a11
almost 70<lspccics. Nowadays. modern authors <Ire more concerned with rcvil'wing
groups than With describing <Ill extensive <lll10Uilt of t<IXCI. l ~xce ption s to this p<llll't"ll
<1re Jochen Martens (Figure 1.1hl cmd Manuel Conzcilcz-Sponga ~Figure 1.1i l. who
h<tvc been working with the rkh faunas of the Himalayas and Venezuela. rcspl't"-
tively. T he tropical faum1s of i\fril'a. i\sia. cmd Centra l South i\nll.:ril"<l probably con-
tfli n <1 large unknown diversity yl'lto be dcsnibcd. nwin!y inthc 111nst diverse l'cuni-
[ies Sclerosomatidae. t:onyleptidat. ;md Cosnll'tidat:. 1\ linute !iJnns. such <IS the
;.:al mox idsa nd Cyphoph thalmi.may il lso providt many mw species. so the re<1 l rkh-
ness of the order may exceed I ~l.tlt)(J species.
Hilrvcst nwn a rc divided into four suborders that cmlt<Jin -+ 5 rl'l'tl)!tlized fa111ilil's
and about I. 500 genera. However. the limit s and rd;rtiunships of most familil's and
genera c1rc imprecise. and nt:w fcnn ilit:s a rc l'XPt'l"tcd to be discowrcd. Clwptcr-+ prt-
sen ts <1 brief morphulogiccd description of cd! supnlgl'lll'l"il' groups. induding l'O IIl-
mcnt s on their distribution. rdationships. main rdi.Tl'lll"l'S. iiiHI kcys. lkprl'St'lltil-
tives of Cyphophtha lmi. the oldest suborder. which indudes (l fc11nilit's cmd 1 ;n
species. ;m.: distributld worldwide. inhe~hiting cd l nmtimnts t~m! isl<llllls of l'llllti-
nental origin. They arc illllOng till' smc dlest :~nd most obsntrc Opilioncs. typiccd(y
measuring between I c11ul ~ mm ill body lcnl!,th. 'l'lll' suborder 1-:upnoi indudl'S h
families and I./X{) spcdt'S. and some Spl'l"it'S o[" th is )-ll"!llljl ill"l' <111101\g tht hl":-.1-
known Opi liunes. They <l l"l' widely distrihutnl in both IHmisph l'l"{'.~. <tnd till' )-ll"l'<ll
111<1jorit y of the Sl}l'des arc sort bodied and loll ).!. lq.!!-!-l'd . Thl' IIWmhns nf Dys p11ui
present a great lliversit y of sir.ts ;md morphologies. i n dudin~ tht' lmgtst lwr-
vestman spedcs. Tm!fu/u .~ torosus. with a body l l Ill In long. This su hordl'r is diddl'd
into 7 f< llnil ies co nt aining 2lJO splties . whkh a rc 11Wi11ly fm11HI i11 the Northem
Hem isphere. l.aniatort'S is the 1nost di\'t'I"Sl' s uhonltr. t'otnpo:-.td of 2h f<llnilil's a nd
t74H SJX:des distribu ted tmtin ly in tropkal atHIIt'llliWr<ttt' rl'gitms 11f till' Stlltllll'l"ll
l lcmisphere. Many rcprescntcti vcs ran rct~l'h l;,rge sizts. Sltl'h <~S 1he gtmykpt id ,\/i-
6 Wllilt Are Harvcstrnerf'

~ :-ooo
~ dXXl

c 5000

l -ID:>O
E
E }();)()
2000

1000
0 ~------~~~=--------------
50 oo :o soaooo t o 20 JO-illSO o0 7oso QQ oo 1o 20 30 4050 60 70 so Q()
LOO ISOO 1900
Decc:ul es
400

10
350
3()()

,
..0
250

,~
,00

ISO
0 100
~ so
t3
m -10 n.:zo 21-30 31-40 41-50 51-60 61-70 71-SO Sl-90 91-00
Decad es

Figure 1.1. (a) Cumulative number of described harvestman species (including only valid
names of living species) in each decade from 1758 to 1999. Note that the point of in
flexion of the curve is in the 1950s, which coincides with the ending of Roewer's career.
The number of species described per decade is strongly correlated with the number of ac-
tive taxonomists (R 2 = 0.637; F:: 40.364; p < 0.001). (b) Cumulative number of articles on
harvestman biology published from 1901 to 20Cl0. It includes only articles (not theses or
congress abstracts) published in English, French, Italian, German, japanese, Portuguese, and
Spanish. Taxonomical works that present only anecdotal information on natural history
were not included in the count. The contribution of researchers from each continent is as
follows: North America, 23% of all articles: Central America, 3%; South America, 18%; Eu-
rope. 47%;Asia, 1%;Africa. 1%; Oceania, 2%
 What Are Ha!Vestmen? 7

Figure 1.2. The ten opilionotogists who have described the most spec1es. (a) The Danish Wil!iam
S0rensen (1848-1916): 157 valid species described. (b) The German Carl Friedrich Roewer (1881 -
1963): 2,260 valid species described. (c) The Brazilian Gindido Firrnino Mello-Leitito { 1886-1948): 347
valid species described. {d) The South African Reginald Frederick Lawrence {1897- 1987): 224 valid
species described. {e) The American couple Marie lOUise Goodnight (1916- 1998) and Clarence J
Goodnight {1914-1988): 313 valid species described. (f) The Estonian, naturalized Braz1han Helia Eller
Monteiro Soares (1923- 1999): 169 valid species described. (g) The japanese Seisho S uzu~1 ( 1914- )
201 valid species described. (h) The German Jochen Martens ( 1941 - ): 147 valid spec1es described. (i)
The Venezuelan Manuel Gonzillez-Sponga (1929-): 219 valid sp~Kies ckscribed
8 Wh<H Arc HilrVL'Strllell?

/r!/'lurslriw r!JIIflls tlcg IV r;rn rL'<.Kh I x:; rnrn). and some of them arc colorful ;r nd/or
1\'dl ;wnwd ll'ith spines
.\ltlwugh tlw liJLrr suborders ;rn: I'L'Cn).!_nizcd ;rs monuphylctk groups, th e rcla
tinnship ;n nun!!, them is sti ll contru\'crsia l (sec Chapter 1). There is no doubt th at
l'yphoph tlwlmi was till' !irs\ group to deri ve. but rnorphologiccrl and molecular
studks IHI\'t' n..nn tly dcrnunstrcr ted that the Dyspnoi cou ld be grouped either with
l: uprHri or l.ani;rh~rts. Hdationships among supt:rfa rnilics/ fmnilies arc also obscure
lirr most groups. nminly the highly divtrst L;rniatures.l\!ost lnrrvestmen. especially
the tropk;r llirrms. ha\'l' ;r l uwd i s pr.: rsalc;~p;~bil i t y and high endemicity. and thus they
urc good rnodels for biogeographic studil's (sec Chapt er 3). Biogeognrphic studies
ll'i!h harn-strnL'll have just beg un. bu t thL'Y may play an important role in our
knm\lcdgt of the L'\ent s that h11ve caused the diversilkation of biotas on both local
il lld rq:.ion al sndes
' ' ' thL onlimrllcwl. h;:rr\'cstmcn arc ubiquitous and c;m be fou nd in a ll cor11i
nents e:-.:n.pt ,\nt;rrt'linr. !'rom the equa tor up to high lcrtitudes (sec Chapter 7J. We
n rn lind tlll'lll in ;1 great v;rril'ty of h<rbil;rt s in <rllterrestrial ecosystem s. includin g in
soil. moss. and k<tf Ii ll er. under rocks. stones, and debris. on vertical s urfaces from
tret trun ks to stone walls. <rmon g grassy dumps. and running over high vegetation .
The most lli\'crsc h;rrvcs trn;rn communities. however. arc reported for tropical areas.
L'S pedally win forests. ,\[t hough some species arc widely distribu ted and ca n be
found in ;r wi de range of habit;,tt s. rmmy harvestman species arc much more limited
in geographic distribution am\ habi t ill USC.L'Spcci ally in tropical areas. Some species
;rre l"L'S iril'tcd to caves. ;rnd others occur in very specllic microhabitats. such as nests
of lcal:.nrl\er ;rnts. The in lluc ncc of physkal factors on the spatial distribution or
lwrwstmL'Il h<rs been poorly s tud ied. but tempera ture and humid ity seem to be the
must import<mt determinants of thei r distribution and habitat use. Despite theca
parity of maintaining in ternal homeostasis. there is s trong evidence that many har
vest man spclies arc inel'1icicnt in avoiding water loss . Some morphologica l and phys
iological features of the order. such as a large surface/volume ratio. lack of spiracular
m nt rol. <md low osmotic hemolymph concentration. may partially explain why most
species arc found in damp nnd shaded areas (sec Chapt er 14).
Phenologica l patterns present marked variation among species (or even popu la
lions of the same species). and few could be regarded as nonseasonal (see Chapter 7).
While most representatives of the suborders Cyphophthalmi and Laniatores arc ac
ti\'e durin g the whole yea r. genera lly with mon thly Ouct ual.ions in population size.
the occurrence of adu lt individuals in representatives of the suborders Eupnoi and
Dyspnoi is usually restricted to a few months of the year. As seen earlier. many har
\'Cstman common names in European countries indeed make referen ce to the phe
nology or the most common species. wherein the occurrence of adults and the
mating season coincides with the harvest period. that is. spring and summer. In tern
pcratc regions some annual species overwin ter as nymphs from eggs hatched in the
r.,ll.,r. ..ui"'<.'J'1..1f&lii d.b t.jrfl.!.li.'i.b>JJr.hr.din. J.hr.:~:;pra,!!.":''111e'ttuhill\:;f'ruraynfy?J.r,1 3n-..
varies a mon g different species. ranging from fou r to eight. but usually six (see
Chapter 13).
 What Are Harvestmen? 9

Most harvestman species have an omnivorous diet that includes small. soft-
skinned art hropods and other invertebrates. as well as carrion. plants. a nd fungi (sec
Chapter 8). This broad a limenta ry spectrum may be considered a unique fct1 turc
among arachnids, whic h are genera lly viewed as exclusive predators of inverte-
brates (mainly arthropods) a nd small ver tebrates. There are a few species th at spe-
cialize almost exclusively in terrestrial s n ails and slugs and thus have t he common
nameSclmeckenkanker, which means "snail harvestman" in German . In order to tin d
food. most harvestman species seem to rely on an ambus h strategy or. more rarely.
on active h un ting. Periods of waiting a re generally in tercalated with periods of slow
movemen t in wh ich the individuals. which a rc unable to form images and probably
c<m only distin guish ligh t from dark. explore their env ironme nt usi ng the lips of
their second pair of sensorial legs. This particu lar feature of harvestman biology in-
spired their popu lar name in Japan. wtomusl ri : zalo is an <lllCicnt \\'Ord used to desig-
nate the players of the bilva. a traditiomll Japan ese stri nged instrument that was gen-
crn lty played by blind people. and muslli means "bug." Therefore. the name
associates the walki ng behavior of harvestmen. with their long sewnd pair of legs
stretching ahead like arms. with a blind person fumbling a long his way with a
wa lking stick.
Unlike most arach n ids. harvestmen do not have <1 pumping stomach to Sllrk the
liquefied tissues of their prey. and they mastkatl' their food by in gesting small
particles (see Chapter 2 ). 1'hereforc. t hey tlrl'l'XI)Osed to paras ites and p;llhogens tllilt
other wise would be filtered out by the feedin g mech;_mism. ilS occurs in the other
arachnids (sec Chapter 9). Grcgarincs are p;1rtirularly abundant in hanestmt'll but
are uncommon in other arachnids. s uch as spiders. Additionally. the omnivorous
feeding habits of many harvestmen place them in dosl' prox imity to cont;un in<lted
mater ia ls. which may resu lt in contact with p;~thogcns or infect in> S t<~ges of some
parasites. These arc not the only natura l encmits of harvestmen: a compihllion uf
the lit era ture clea rly s hows that the most di w rsc group of har\'l'stman pred:1tors is
that of passerine birds(seeChapter 9). Fwgsand touds. in strtivtlnJu s lll<lmln<lls. <lnd
spiders nrc oth er importc1nt groups of predc1tnrs
'lb dea l with this diversilied ran ge of natura l e nemil's. lmrwstmt n haw dl'\'l'l-
oped a great variety of dcfensi\'l' stratc:gies (Sl'l' l'h<iptl'l' H)l. Sonw spt.'l'il's. fo r
insta nce. cmnouflage th emselves with debris glued on hy il St'l'l'l'liml from tlw in -
tegume nt. and very frequently they respond to pred;1tor a \larks hy t'c:igning dtc 1\h. a
defe ns ive behavior known as tlmnatosis (al'tc r the l:rlck god of dl'il t h. TIHHl<lt osl. If
disturbed. many lon g-legged species n1 p idly vihn1tc tlw body. a defcnsiw lwiW\'ior
known as "bobbing" th <ll prob;1hly nmfuscs till' identilkntion <IIH! L'Xtll'l lm;1tion ol
the harvestman's body. Because of this last defense mt-r h<~nism. i11 SOlllL' pl<tl'L'S ol
southeastern Brazil the long-legged IHif\'Cslman stxdes il l'l' ca lled llntnlms-llltiluriwts.
wul in Cos1<1 Rica th ey ill'e known <IS /WIIIIIjos !Tc1hle 1.11.
t\ s we h;1vc seen earlier. nu1ny l'lllllllHlll lltlllll'.'i till rihlltLd to lw rwst llll'll <lfl!liiHI
the world arc rdat ed tot he bad smell nr
thl'ir snnt glands ;uul to t lwir kg lllnw-
mcn ts afier they arc shed . These two dcllnsin stn1 tq.:ks art so conspicuuu.' t h;11
they ht~ve eve n a ttracted the att en tion of IHIIll'.x pcrts in !Ill' grt!llp. such :~s tiH' 1';1-
1Q Wh,\1 AIC' H,HVC'SI IllCtl 7

111ou:-. Sp:111i:-.h painll'r Sal\';~ d nr Dali. whose inltresl in bizarre animals is (juilc cv i ~
dc111 i11 lib 1\ ' Jlrk ~. l1 1 hi:-. I 'J-1! 1 p;tinling nr~dd!!l,JJII.i/ll'.f/.~ ~~ llw t:l'f'lliny- 1/opl'l Dedi
,;~rdully illuslfiill'lllwonu nnln ll ddtnSt slral q.til's in Opiliunts IFigurt 1. ~). Tlw
harw:-.lman. dc;uly il nprt'Sl'lll<lliwuf l lw s uh,,rdlT Eupnoi. IK'l'Upics a t:cn tral pnsi-
lillll in 1h c p:1i1lling 0111d h:~ s h1s l lllll' uf il s legs. 11 deknsiw str<1\l.:gy known <I S ii U-
IIl~ p<~:-.y I"<T l'h<~pi <T I ill. i\ liWt'onr. the individu al is sur rounded by a swarm of ant s
1lwt l'<~illunull:u:l tlw hantstnl<lns kgs . proh:~b ly IK"cau se of the m:lion of sccnt -
~l<~nd snnlillll S. \\'hil'l1 :1n' knll\1'11111 ht a highly dfectiw repellent <~gain s ! th ese in-
..,,.,ts . . \llhou gh IIll' a 11nosphcn: uf I he p<~i nti n).! is hb1k.thc han'l.'Si nwn i ~ a rc!Cr-
l' IH'l' 111 ill I 11ld Frl'Jldl Pl'iiSillll lq!l'lld 111<~1 s; 1ys !hilt the si!:( llling of a harwstman in

Figure 1.3. The painting DaddyLongfegs of the Evening-Hope! (1940, oil on canvas. 10 x 20 inches) by Salvador
Dali, which is an allusion to World War II. A grieving cupid appears in the lower left of the painting. tormented be-
cause the world is violently changing. In the center of the painting is the daddy longlegs, which, according to an
old french legend, is a symbol of good luck. Thus, in spite of the bleak atmosphere of the painting. Dall offers
hope. Reproduced from Collection of Salvador Dali Museum, St. Petersburg, Florida: 0 2006 Gala-Salvador Datr
Fundation, Figueres {Artists Rights Society. New York): 0 2006 Salvador Dall Museum Inc
 What Are Harvestmen? 11

the evening hours is a good ome n. a portent of good luck. and a sy mbol of hope. In-
terestingly. the harvestman's common n ame in Turkish ( miijtl11'i or miictlll'i) suggests
a similar symbolism. since it means a person who brings good news (Table I. I ).
Unlike ot he r arachnid groups. ma ny species of Opiliones seem to be highly tol-
erant of conspccifics. Several species form dense diurnal aggregations consisting
mainly of subadu lts a nd adults of both sexes (see Chapter I I). In general. ind ivid-
u;:~ls aggregate a t protected sites and close to a wa te r source. The number of individ-
uals in these groups ranges from 3 to nearly 200 mnong the La niatorcs. but a mo ng
the Eupnoi there arc records of mass aggregations conta ining more than 70.000 in -
dividuals. One of their common names in the United States. "grandfat her-gray-
beard ." is probably related to these hu ge aggregati ons since ind iv iduuls ure ft1cing
upwa rd and their long legs arc hanging down. resemblin g a beard or wig. Al though
gregariousness in harvestmen seems to be primarily indun.'<l by env ironmental fm-
tors. this behavior may confer severa l defens ive advantages. in cl uding strt n~th
e ning of defensive s ignals through the collective release or scen t-gl tmd secretions.
speedin g the response to alarm signals provided by the srent set-rt!ion. <1nd de-
creasi ng each individual's chances of being eaten !dilution dtlrt )
The great majority of harvestmen reprodun sexually. although sonw species re-
produce asexually by parthenogcntsis. an d tht Sl'X chromosomes hm'l' been idellli-
lied <IS usually XV-XX (sec Chtlpter h) . As Wl' saw c<~ rli cr. h;~rvl'SlnKn nwy lwvc beLn
the first group of arthropods to t'\'olvc an intromillent org<lll. this bl'ing <IIWtlwr
unique morphologin1 l feature of tlw order a mong arach n ids. llowt'\'L'r. tht study of
mating strategies is probably the 11spect o f their behavior that ha s reniwd thtlt;~st
atttn ti on . Harvestman fcrtili.-:ation is intcrnt1l. and the lr<lllSflr 111" spcrlll< lltiZtli l lll<IY
occ ur indirectly throug h spernwt ophon:s (in Cyphophtha lm i) or dirL"L"tly by 11\l'tlllS
of long and fully intromi ltent mal e genita lia in the ot her su horders ISt'l' Clwpll'r 121.
Unlike courts h ip in other arachnids-a pronss in whidl malts 1\HISt lirst per-
suade" the fema le not to considtr them a s prey ;md th<JI may require <ll'<lrt'ful ap-
pnmch and a long-di sltlnce. chlhnn ltcd. highly stcrLotyptd. <llld spccil's-sperilk \'i-
sua l or vibratory set or displtlys-courts h ip bdurc intnun issiun is ulkn quirk and
I<Jl'tile in har vest men . In some cases. hmn:wr. males m;1y tJI"!i.r a gla1Jdulo1r seLTl'litJn
of their c hcliccr;Jc bdi1re ropu laliun as a nupti;1l ;.tift ti1r their mall'S. 1\1<1ny swdiL'S
a lso mention int e nse courtshi p dt1ring i111 nJnJi ssitJJI t111d m;tlt" gt1arding <ll"ltr npt lltl -
tion. i\dditiomdly, males of many spel"it'.'i dl'l(.>nd territories. ll'hich t~rv used by li.-
m<t les as ovipo:-;ition si tes. Ciwn the ;una.-:i11g eorllpll':dly of tile rmllt ;.:tnitalit~ <111d
the enormous diversity or typts of sexual dimorphism. sex ual Stkdion !ht it imra-
or interscxutd) has probably pl<~yetl a m<1j1 Jr rt1le in the en1lut itrn uf lwrwstmen
Females m<IY lay th eir c~gs imml" di <~tdy ur IIlii Ill hs al"ltr nlpult lt i1111. ;ard 1hLlggs
may ltrkc from 111 days to mort lh<~n livt IJHIIllhs to h:~t rh \Sl'l' l'ht~plcr l ll . Thl
forms or pa rentaleare may indude the pnxluction of l<rf).!l' yolky eggs. till" prtparil-
tion or nests. the c hoice of 01 ppropriall' ovipw:it ion sill"~. ;u1d piirLntal care \Sl"t'
Cha pter 11). Alt hough maternalc<lrl' is wi{ksprc<~d <UJHIII).! !lrirl"h11ids. l~:lr w~lllll"ll
a rc the on ly order in wh ic h some spl:rks prescntexdu siw P<l lt"l'll<il Cilrt'. till r<~rl..;t
form of paren t;d investment ;rn1ong ;rrth ropods. Both lll<llt'ntid ;md p<ill"l'll;il Ciln
12 WIWt AIC HmvestlllCil?

haw lwln dl'llHJtlSir<ltl'd to pl:1y 01 crud;il role in egg su rviva l. preventing prcd<J tion
0111d rull!-!i \l ll al'k . 111 11\'l'\'er. tl ll' :-;delti W rlll"l'CS k<iding I ll th e evol ution of these tWO
lili"IHS of P<ll'l'llt<l assistciiH'l' St'l'lll to be w ry d ifferent. Sin<:e the grc;~ t majorit y of
lll<lil'rll<il lwr ws tllll'll <ll'l' l'l'Stril'tcd to '' si ngle reprodur.:livc event during the
hrccdi11g St'O IStu l. ftnwks l'illl ad1il'\'l' grccl ttr rcprtKiu ctivc success by remai11ing
duse to their holll'hlings throu g hout the l'ari ng pcritKl. Thus ma tcnw l ca rc is likely
to Jww en1hed <IS <1 resul t of na tured sdect ion. In contr<Jst. male t'arc in harves tmen
Sl'l' lns In lw\'l' evolwd <ts <1 reslilt of stx ual sdeltion . i\n:ordin g to ret.:ent studies.
m:1ks th<1t provide pattrnal rare <ll't' preferred by rem:1ks and obtain a g rc;~tcr
11\llllbl.'r of ropui<J tions tlw n nwks th;~t ;1 rl.' llililbk and/or un willi ng to provide ca re
r\lllwugh h:,rwstllll'tl arc 11 f:J scilw tin g group of ;~r:~ r hnid s. th e dram~tl k' in-
l'rl'iiSt' in tmimnnwntal disturba nce <roUIHI the world--especially in tropic<tl rc-
gions- nwy hiiW driwtllll<lllY species to extinction even before their for ma l descrip-
tion hy tc1:wnomists. 1\ lcmy huma n acti vities. includin g pesticide usc. forestry
operations. ilir <l!ld soi l poll u tion. lire. and even th e introduction of domestic an i-
mals.IW\'c trt'llll'llllou s impart:-; o n the habitats on wh ich hilrvcstmen arc dependent
tSt'l' t'lwpttr 71. !\ lost harvest man sptdes have restricted distributions and can be
pa rtit'ulculy end<1ngcred ir human <ll'l ivitil's unfavorably a lter their habitat s. Thi s is
tht case fu r most caw dwclkrs. wh kh may be drii'Cilto l<Kal or complete exti nction
if th e cc1vern irolous h;ibitat ur th e nearby ex ternal environmen t is severely dis-
turbed. In fm l. almost al l lwrvc.stman species form ally considered endangered
around the world ilre cave dwellers. although many other species li ving in other
habit;1 ts may he C<Jtlillly tnd:m gertll. Ho\\'evcr. given the paucity of ecologica l infor-
nwtion for the greil t majorit y of IHirVL'St man species. it is almost impossible to rcli-
<lbly report on their conservation s t <~tus. Th erefore. the preservation of habit ats in-
StLad of partku lar SIX'l'ics may be the most effective mea ns to protect the diversity of
Opilioncs around th e world .
Giwn their geographic distribution and species richness, it is surprisi ng tha t har-
vest men have remained largely ignored by the genera l public and zoologists as well.
Their bthavior and ewlogy arc just beginning to be understood. and in the last two
decades the number of studies published on these subjects has increased consider-
ably, especially in South America (Figure 1. 1b). Populations of many species arc lo-
cally abundan t. and individuals arc easy to observe. Chapter 1 5 providt.'S a n
overview or th e methods and techniques used to study harvestman biology both in
tht Jicld and in the laboratory. There you will learn how to collect specimens for eco-
logical samplin g. prepare material for cytogenet ic studies. preserve pathogens and
parasites of harvest men , and conduct taxonom ical and systematic studies. More-
over. you wil l discover that representatives of the order may be easily maintained in
taptivity, where they be/Hive in a similar way to that observed in the field . In conclu-
sion. harvestmen arc perfect subjects for ecological, bch<1vioral. and evo lu tion <~ry
studies. and we truly expect I he reader-whether an am ateur or a professional- to
li nd the book a usefu l tool for progressing in the understanding of the fascinat ing bi-
ology of harvestmen.
 What Are Harvestmen? 13

ACKNOWLEDGMENTS

We arc grateful to those who helped with popu lar harvestman names: Astri Leroy
and Adriano B. Kury (Afrikaans ). Ch ristian Lcxcr and r:.cti th Karpinos (German).
Gabriela Lysakova (Czech). Gilbert Barrantcs (Spanish-Costa Rica), Grace Young
Kim (Korean), joa nn a Znaniecka (Polish ). Laszlo Csiba (Hu ngarian). Luc Van hcrckc
(Dutch). Luis E. Acosta (Spanish-Argentina). Luis l nda (Spanis h-Spai n ). Marcc ltl
VanLoo (Slovakian). Michael Fay (Welsh ). Niklas Wahlberg (Finn ish and Swedish ).
Nobuo Tsurusaki UapH.nesc). PI amen M itov (Russian). Shan Luan (Ch inese). Sorcn
'Jbft andThco Blick (Danish and Norwegian). Tone Novak (Slovcnian). iJ mit Kcbtlpd
(1\lrkish ). Wojcicch Starcga (German. Polish . and Ukrain in n ). and Sa 11Hm th a
Koehler. who helped us cont;,tct most of these people. Ron Clouse ki ndly nviscd this
chapter. We arc deeply indcbtL>d to Bruno A. Buzatto fo r helping assemble the litera~
lure fo r Figure 1.1 b. to Joan R. Kropf and Carol Butler for givin~ us permission tore-
produce the painting Dadil!l l.ounkys of till' EI'CIIiii!I-Hopc! ;md the following m l-
lcagucs for providing the photos in Figure 1.2: Adriano It Kury. Ann Bndwnowski
Nikolaj Sc harfr. Nobuo Tsurusaki. Peter Jiigcr. and Zandi k ~lbhdc