Table Of Content1993. TheJournal ofArachnology 21:120-146
PATHOGENS AND PARASITES OF OPILIONES
(ARTHROPODA: ARACHNIDA)
James C. Cokendolpher': Adjunct Professor, Department ofBiology, Midwestern
State University, Wichita Falls, Texas 76308 USA.
ABSTRACT. ThisisthefirstpapertoreviewtheliteraturerecordsonallpathogensandparasitesofOpiliones
on a global level. These organisms (bacteria, fungi, protozoans, cestodes, trematodes, nematodes, arthropods)
are listed in phylogeneticorderalongwith available information on hosts, collection localities, life history, and
taxonomic history. The opilion hosts are also listed (by their currently accepted names) along with the names
oftheirknown pathogensand parasites. Diagnosticcharacters and some taxonomic keysare providedfortaxa
whichare relativelywell know. Citationstootheravailablekeysare provided. Manynewhostanddistribution
records are provided.
Two fungi [Engyodontium aranearum (Cavara), Torrubiella pulvinata Mains] are removed from the list of
pathogens ofopilions and it is suggested that the original hosts were misidentified spiders.
Two new combinations are recorded in the Mermithidae: Agamomermisphalangii (Haldeman 1851), Aga-
momermis truncatula (Rudolphi 1819). Agamermis incerta Steiner in Stipperger 1928 is regarded as a nomen
nudum.
The type locality ofthe mite Leptus lomani (Oudemans 1903b) is restricted to Corral (39°53'S, 73°25'W),
Valdivia, Chile.
Unlikemanyarachnids, Opilionesorharvest- temal pathogens and parasites (see Holmberg
men lackapumpingstomachandthereforethey 1986, and citations therein). To date, only de-
chew their food and often consume oocysts and fense against predators and harvestman aggre-
spores. Examination oftheir feces reveals a va- gation formation have been demonstrated.
rietyofchitinousfragmentsfromtheirarthropod While working with a South America har-
prey as well as plant pieces. Some saprophytic vestman, Estable et al. (1955) discovered that
fungi and yeast spores can be observed as well the exocrine gland secretion was a remarkably
as gametocytes of internal parasites. The fre- effectiveantibiotic, in vitro, against 18 genera of
quent grooming of the legs by the harvestmen bacteria (Gram positive and negative) and pro-
may also lead to the ingestion of oocysts and tozoa.Theirworkrevealedthatthesecretionwas
spores. While ingestion is the common entrance also active when given orally to mice infected
pathwayforsomeopilionpathogens,fungiinfect with intestinal parasites. The substance was tol-
theirhostthrough penetrationofthe cuticle. Al- erated perfectly by the mice but destroyed giar-
though gregarines and mites are frequently en- dias, trichomonas and hexamites. The compo-
counteredwhen observingharvestmen, relative- nents ofthe secretion were later determined to
ly few researchers have documented their be a composed ofa variety ofquinones (Fieser
occurrences. & Ardao 1956).
Harvestmen are unique amongarthropodsby The major components ofthe exocrine secre-
possessing bilateral exocrine glands which open tions ofharvestmen differbetween the two sub-
ontothedorsalsurfaceofthecephalothoraxnear orders, Laniatores and Cyphopalpatores. Minor
the base ofthe second pair oflegs. These glands components and ratios of components differ
produce a variety ofvolatile secretions (Ekpa et amongcongenericspecies(Ekpaetal. 1985).The
al 1984, 1985) thathave beengenerallyconsid- fewchemicalanalysesthusfarreported(seeEkpa
ered to be defensive in nature. The glands have et al. 1984, and citations therein) from Lania-
also been proposed to function in a variety of tores reveal a variety of alkylated benzoqui-
other behaviors including protection from ex- nones, phenols, N,N-dimethyl-i(?-phenylethyl-
amine and bomyl esters. Only the Palpatores
'Home address: 2007 29th Street, Lubbock, Texas sectionofthesuborderCyphopalpatoreshasbeen
79411 USA. chemically investigated. Those analyses reveal
120
COKENDOLPHER^PATHOGENSAND PARASITES OFOPILIONES 121
membersofthisgroupsecreteshort-chainedacy- topicofphoresyarises. Phoresyisnotparasitism
clicketones, alcohols and naphthoquinones (see but rather a form of symbiotic relationship in
Ekpa et al. 1985, and citations therein). which the smaller organism associates with the
Eventhoughharvestmenareabundantinwarm harvestman in order to obtain transportation.
moistsituations,fewrecordsareavailableoffun- Phoresy as well as passive transport of fungal
gi infecting these animals. Because the major and plant spores will not be examined here.
components ofharvestman exocrine secretions
are members of chemical classes known to be
Superkingdom Prokaryotae
fungicides(seeTorgeson 1969; Coleetal. 1975),
Kingdom Monera
it is likely these secretions are used to protect
harvestmen from infection. The use ofthese se- Division Gracilicutes
cretions in defense and grooming needs further Family Enterobacteriaceae
study. Xenorhabdus Thomas & Poinar contains five
Whilethereareseveralworld-widetaxonomic described species and other undescribed species
revisionsofharvestmen,nosimilartreatmentfor (Akhurst & Boemare 1990). They typically in-
their parasites has been undertaken. This is in habit nematodes and their host arthropods (in-
part due to the incorrect view that harvestmen sects and arachnids). See under Nematoda
are not ofeconomic importance. Mounting ev- (Rhabditoidea) for further details on this rela-
idence demonstrates that harvestmen are bene- tionship. Pertinent taxonomic papers are cited
ficial and that they consume considerable quan- with areviewofthetaxonomicproblemsinAk-
tities ofpest insects and mites. Because ofthis hurst & Boemare (1990).
beneficial status, no one has investigated para- Xenorhabdus luminescens Thomas & Poinar
sites for controlling opilions. Experiments in- (1979) is introduced into the arthropod host by
volving insect pathogens on harvestmen reveal a Heterorhabditidae [Heterorhabditis bacterio-
opilions are susceptible. Like conventional in- phora Poinar]. Poinar & Thomas (1985) dem-
secticides, insect pathogens and parasites could onstrated this bacterium could kill a Phalangi-
have a severe impact on the beneficial harvest- idae{PhalangiumopilioLinn.,reportedasP.sp.)
men. ifintroduced by the correct nematode.
Many of the records of parasites from har- Xenorhabdusnematophilus(Poinar&Thomas
vestmenareincomplete. In somecasesthe host, 1965) was originally described in combination
but not the parasite, is identified to species. In with Achromobacter Bergey, Breed & Murray.
othercases, the parasitebutnotthehostisiden- Thisbacterium is introducedinto the arthropod
tifiedtospecies.Thepurposeofthiscontribution host by a Steinemematidae [Steinernema car-
is to bring together the limited information on pocapsae (Weiser)]. Poinar & Thomas (1985)
this topic so that a foundation can be built for demonstrated that this bacterium could kill a
future research. Phalangiidae, Phalangium opilio (reported as P.
Becauseofthelackofgoodcharactersinsome sp.), ifintroduced by the proper nematode.
groups(/. e., MicrosporidaandjuvenilesofMer-
mithidae) collective groups have been named.
Superkingdom Eukaryotae
Suchgroupsorgeneraoftenincludespecieswhich
probablyarenotrelated.Thisgroupnameisused Kingdom Fungi
Division Eumycota
simply for “taxonomic convenience” and in-
cludesspeciesnotreadilyplacedinknowngenera Atleastonespeciesoffungussuccessfullykills
(possibly because a particular life stage is un- a Gonyleptidae (see under Torrubiella gonylep-
known) and species incertae sedis. Some taxo- ticidaandunidentifiedfungi).Gonyleptoideaare
nomicallyconvenientgroupsalsooccurathigher known to have phenols which are antagonistic
levels in fungi. In fungi, the sexual stage (teleo- to fungalgrowth intheirexocrine secretions. Ei-
morph ofascomycetes and basidiomycetes) and ther T. gonylepticida and another unidentified
theirasexualstages(anamorphsorconidialstages) fungusfromPanamaarenotretardedbyphenols,
are sometimes placed in separate genera and or the hosts were unable to produce phenols in
classes. In somecases, two ormoreascomycetes sufficient quantity. The extent of phenol pro-
may be identified as having the same form spe- duction in variousgonyleptidgenera and its use
cies for an anamorph. in controlling fungi have not been investigated.
When one discusses parasites ofopilions, the Likewise,theeffectsofageandhealthofthehar-
122 THEJOURNALOFARACHNOLOGY
vestman on phenol production have not been Spicaria longipes; which is now recognized as
examined. Paecilomyces farinosus (Holm ex S. F. Gray)
(Brown & Smith 1957), Fetch recorded T. gon-
yleptidda and the conidial stage from various
Unidentified Fungi spiders from Trinidad. KovaF (1974) listed the
Griffiths (1978) illustrated a harvestman [not conidial stage from spiders collected on Mag-
identified, but almost certainly Nelimapaessleri nolia Linne leaves in Russia (formerly Russian
(Roewer)]coveredbymyceliaofsoilmicrofungi. Soviet Federative Socialist Republic, USSR). In
The fungiare reportednotto be pathogenic, but the key to Torrubiella spp. by KovaF, two va-
simplyusetheharvestmancorpseasasubstrate. rieties of r. gonylepticida are differentiated on
Mora(1987, fig. 8)reportedmortalityinadult the basis ofperithecia and ascus lengths. How-
males ofa Gonyleptidae {Zygopachylus aibom- ever,thetwotaxashouldbeattributedtoanother
arginis Chamberlin) by an unidentified fungus species: the third taxa in the key should be T.
onBarroColoradoIsland,Panama.Malesofthis arachnophilavar.pleiopusMainsandthefourth
nest-building harvestman eat all fungus appear- should be T. arachnophila var. pulchra Mains.
ing in the nest, thus preventingthe proliferation Torrubiella pulvinata Mains (1949) was de-
ofmycelia. Mora (1987) suggested the males in- scribedfrom “Opilionoidea”collectedonOahu,
gested the fungi (spores) which eventually killed Hawaii. Paecilomyces(reportedasSpicaria)pul-
them. This is probably incorrect because nearly vinata (Mains 1949) was the name given to the
all otherfungal pathogensofinvertebratesinfect conidialstage. Samson(1974)listedS.pulvinata
theirhostthroughthecuticle(Samsonetal. 1988). as a synonym ofP,farinosus, thus regardingthe
Ten fatalities were observed from 199 nest- anamorph for both T. gonylepticida and T, pul-
guarding males examined by Mora. vinata to be the same species. Mains (1949, p.
303) stated “The hosts ofthis collection are so
severely parasitized that accurate determination
Subdivision Ascomycotina is difficult. They appear to be arachnids belong-
Class Pyrenomycetes ing to the Opilionoidea.” Because opilions ap-
Order Clavicipitales pear to be absent from the Hawaiian Islands (F.
Family Clavicitaceae G. Howarth pers. commun.), the host is more
likely a long-legged, pholcid spider. The setae-
Torrubiella Boudier is a genus with primary spinesonthelegsillustratedbyMains(1949, fig.
host affinities for spiders (Araneae), although
lA) are long and unlike those on harvestmen.
several species are also known from insects, es-
pecially Coccidae (Kobayasi & Shimizu 1982; Theyaresimilartothosefoundonspiders.There
Humber & Rombach 1987). Two species have arefiveadventivecosmopolitanFholcidae(Ara-
neae) established in the islands that could be
been reported from harvestmen, but only one
confused as opilionsby non-specialists. The op-
report appears to be valid.
ilion host records are considered here to be in-
Torrubiella gonylepticida (Moller 1901) was
correct.
originally described in combination with Cor-
dycepsFries. Fetch(1937)transferredthespecies
to its present combination and redescribed the
species. Moller(1901),whendescribingthehost, Subdivision Deuteromycotina
referredtoitasaspider(‘DieSpinnen’,not‘We- Class Hyphomycetes
berknechte’). Subsequent authors (Fetch 1937; The Hyphomycetes is an artificial class rep-
Koval 1974; Kobayasi & Shimizu 1982) have resenting the asexual states ofAscomycetes and
continued to list the only host as a spider. For- Basidiomycetes, or fungi forwhich sexual states
tunately,thespecificnamereferstothetruetype are unknown. Orders and families do not exist
host, aGonyleptidaeharvestman. Moller(1901, in current classifications ofthese fungi.
taf. 6, fig. 89) clearly illustrated the gonyleptid HymenostilbeFetch iscomprisedofseven de-
host, but not in sufficient detail to determine to scribed species. Species are known to infect a
which genus it belongs. Kobayasi & Shimizu varietyofinsecthosts,spiders(Mains 1950;Evans
(1982) reprinted MolleCs illustration and stated & Samson 1987) and harvestmen. Mains (1950)
the type locality was Brazil. stated members of this genus are the conidial
Fetch (1937) described the conidial stage as (anamorph) state of Cordyceps spp.; whereas
COKENDOLPHER-™PATHOGENS AND PARASITES OFOPILIONES 123
Evans&Samson(1987)reportedtheteleomorph Subdivision Zygomycotina
connection remains unproven. Specific identifi- Class Zygomycetes
cationsarebestmadebyconsultingthediagnoses Order Entomophthorales
provided by Mains (1950). Hymenostilbe ver- Family Entomophthoraceae
rucosa Mains (1950) was originally described Pandora Humber (1989) is comprised of 16
from spiderscollectedin Maine, USA. Otherre- species ofobligately pathogenic fungi. Hosts in-
cords are from spiders in England and a “Phal- cludemembersofinsectsandarachnids.Asingle
angiidae” in England (Leatherdale 1970). speciesisrecordedfromopilions. Pandoraphal-
Engyodontiumde Hoogiscomprisedofseven angicida (Lagerheim 1898) was originally de-
species, two ofwhich are reported to infect spi- scribed from Phalangiidae collected in Sweden
ders and one on '‘opilionids”. A key to the spe- as a species ofEmpusa Cohn (Entomophthora).
cies is provided by Gams et ah 1984. Engyo- Batko (1966) transferred the species to Zoo-
dontium aranearum (Cavara) was originally phthora Batko 1964, and placed it in the sub-
describedinthegenusSporotrichumLinkexFries genus Pandora Batko 1966. Humber (1989)
and was transferred to its present combination placed it in his new genus Pandora. Ellis (1956)
by Gams et al. (1984). A redescription and syn- and Leatherdale (1958, 1970) recordedthis fun-
onymy are providedby Gams etal. (1984). The gus from a Phalangiidae, Phalangium opilio, in
teleomorphstateisunknown,butothermembers England.
ofthegenushaveaTorruhiellateleomorph.Those Entomophaga Batko includes 10 described
sameauthorsreportedhostsasa fly, spidersand species (Humber 1989). All are obligate patho-
opilions. The specimen in their photograph (fig. gens ofinsects and arachnids. A single species is
3), as well as those ofSamson et al. (1988, pi. recorded from opilions. A key for identification
68a,b),superficiallyresemblesopilions,butjudg- ofmembers ofthis genus is provided by Keller
ing from the dense placement and morphology (1987). Comparisons to original descriptions
ofthe setae on the host legs (figs. 3, 68b) they (speciesandcitationsarelistedinHumber 1989)
arenotharvestmen.Theyaremorelikelypholcid are required for positive identifications. Ento-
spiders (Araneae: Pholcidae). The opilion host mophagabatkoi(Balazy 1978)wasoriginallyde-
record for this species of fungus is considered scribed in the genus Entomophthora Fresenius.
herein incorrect. Later, Remaudiere & Keller (1980) transferred
NomuraeaMaublanciscomposedofthreede- the species to ConidiobolusBrefeld (Family An-
scribed species (Ignoffb et al. 1989; Greenstone cylistaceae), but the current combination with
et al. 1988). Nomuraea rileyi (Farlow) Samson Entomophaga was made by Keller (1987). BaJ-
is a well-known pathogen ofinsects. Nomuraea azy (1978) described this fungus from harvest-
atypicola (Yasudo) Samson is reported to infect men collected near Poznan, Poland. Phalangi-
spiders,harvestmenandinsects.Nomuraeaane- idae[Oligolophustridens(C.L.Koch)]andrarely
monoides Hocking was originally isolated from Sclerosomatidae{LeiobunumrotundumLatreille
soil and, in high doses in the laboratory, can andLeiobunumblackwalliMeade)wereinfected.
cause mortality in insects. An epizootic (temporary increase in the inci-
Nomuraea atypicola (Yasuda 1915) was orig- dence of infections) was observed during late
inallydescribedasamemberofthegenusIsaria summer.
J. Hill exE. M. Fries. It was found on an Atyp- Keller (1987) reported this species of fungus
idae spider in Japan. It was transferred to its was rathercommon and often causedepizootics
presentcombination by Samson (1974). The te- in open woods, along the borders offorests and
leomorph or sexual state is Cordycepscylindrica hedges. From lateJulyto the middle ofSeptem-
Fetch (1937). Greenstone et al. (1988) reported ber it was collected from Oligolophus tridens in
the infection ofa harvestmen bythis fungus un- Switzerland.
der laboratory conditions. The infected Sclero-
somatidae, Leiobunum vittatum (Say), was col-
lected in Missouri, USA. This species offungus Kingdom Animalia
is commonly found infecting spiders (Green- Subkingdom Protozoa
stone et al. 1988) and under laboratory condi- Althoughseldomreported. Protozoaarecom-
tions was also found to be infective to Lepidop- mon parasites ofOpiliones. To date, all records
tera larvae (Ignoffb et al. 1989). ofProtozoanparasitesofOpilionesarefromUSA,
124 THEJOURNALOFARACHNOLOGY
Europe and India. Their reported absence from the feces, and no intermediate host or vector is
otherlocalitiesislikelyduetolackofstudy.While needed. Because mostspecies arebelievedto at-
dissecting gonads for anatomical and chromo- tach to intestinal epithelial cells, gregarines in
somal studies, I have often observed gregarines opilions probably are not pathogenic.
from North and Middle America species (es- Thegregarinegeneraandsomespeciesknown
peciallyfromPhalangiidaeandCosmetidae). El- fromharvestmencanbeidentifiedbythefollow-
lis (1913, p. 280) reported thathe was unable to ing taxonomical key. Because some species are
locategregarinesinthe“alimentarycanalofper- inadequately described (some life-stages un-
haps two hundred Phalangidea” from Michigan known) identifications to species are difficult.
andColorado.Hisfailuretolocateparasitesmay Usefulkeysortablesofcharactersarementioned
have been caused by the time ofyear he exam- under specific genera in the following account.
ined the opilions or possibly the taxa he exam-
ineddonotharborgregarines(thesetaxaareun- Class Sporozoasida
known, but probably are members of the Subclass Gregarinasina
Sclerosomatidae: Leiobuninae as they are the Tsurusaki (1986) found gregarines in Sclero-
dominate forms in the two mentioned areas). somatidae harvestmen, Leiobunum manubria-
Onlytwostudieshavebeenpublishedonopilion turn Karsch and Leiobunum globosum Suzuki,
hematocytes, one ofwhich resulted in the dis- from numerous localities inJapan. He also pro-
covery ofa blood parasite. vided data on parasitism rates as related to spe-
cies, localityandseason. Hisgregarineshavenot
Phylum Microspora
been identified.
Class Microsporea Hunt(1979)foundnumerousgregarinesinthe
Order Microsporida
midgut diverticula ofTriaenonychidae harvest-
Collective Group Microsporidium men, Equitius doriae Simon, from southeastern
Speciesthatcannotbereadilyplacedtogenus, Australia. His gregarines were never identified.
as well as species incertaesedis, are lumped into Mitov reported (pers. commun.) that he had
Microsporidium sensu Sprague (1977). Micro- discovered gregarines in preserved material of
sporidium weiseri (Silhavy 1960) was originally the following harvestmen from Vitosha Moun-
describedinthegenusStempelliaLeger&Hesse tain and West Rodopy, Bulgaria: Nemastoma-
(Family Thelohaniidae). Sprague (1977) trans- tidae [Carinostoma ornatum (Hadzi), Parane-
ferredthisspeciestoitspresentcombinationwith mastoma radewi (Roewer), Pyza bosnica
Microsporidium because the species did not fit (Roewer)]; Sclerosomatidae [Leiobunum rume-
anyoftheknowngenera.Thisparasitewasfound licumSilhavy]; Phalangiidae[Laciniusephippia-
in smear-preparations ofhemolymph ofa Phal- tus(C.L.Koch),L.horridus(Panzer),L.dentiger
angiidae, Opilio parietinus (De Geer). The har- (C. L. Koch), Lophopilio palpinalis (Herbst),
vestmen was collected in Treble, Czechoslova- Mitopus morio (Fabricius), Odiellus lendli (So-
kia. Theplasmodium have 2, 4, 8 and 16 spores rensen), Opilio dinaricus Silhavy, O. ruzickai
andarefoundinthehemolymphandhemocytes Silhavy, O. saxatilis (C. L. Koch), Phalangium
(plasmatocytes) ofits host. opilio, Zacheusanatolicus(Kulczynski), Z. crista
(Brulle)].
Phylum Apicomplexa
Other new records include unidentified greg-
All known Apicomplexa parasites ofopilions arinesfromaPhalangiidae, OdielluspictusWood,
areseptateeugregarinesandassuchhaveseveral collectedintheWestVirginiaUniversityForest,
features in common. Both sexual and asexual ChestnutRidge, Preston County, WestVirginia,
stages occur (gametogony and sporogony), but USA and an unidentifiedgregarine from a Scle-
merogony is absent. The mode of infection is rosomatidae,LeiobunumpolitumWeed,collect-
ingestion ofoocysts. The trophozoites attach to ed in Columbus, Ohio, USA. This latter series
the lining ofthe gut and divide to form mero- is remarkable as the parasites were only discov-
zoites and gamonts. Gametocytes are passed in ered after a hundred years ofstorage.
Key For Identification OfGregarines Found In Harvestmen
1a. Oocysts without spines orthickeningat poles 2
1b. Oocysts with spines orthickenings at poles, sometimesat equatorand also alongedges
(Family Actinocephalidae) 5
.
COKENDOLPHER-PATHOGENSAND PARASITES OFOPILIONES 125
SubfamilyAcanthosporinae
2a. Epimerite simple, spherical; oocysts biconical, with truncate ends, released unchained by simple de-
hiscense ofthe gametocyst Family Hirmocystidae
Arachnocystisarachnoidea(Devdhar& Gourishankar)
2b. Epimeritecomplex and varied; oocysts biconical orcylindroconical, united as a stringofbeads
FamilyActinocephalidae 3
SubfamilyActinocephalinae
3a. Epimeritesessile, with shortneckhaving8-10 simpledigitform processesatapex; neckpersistsmore
orless in sporont, butdigitform processes (tentacles)disappear; gametocystsdehisce by formation of
hole in wall throughwhich oocysts are extrudedin a single thread; oocysts biconical orlemon-
shaped Actinocephalus megabuniOrmieres & Baudoin
3b. Epimeritewithout digitform process atapex, gametocystsrupture by simple dehiscence 4
4a. Epimeritea large, flattenedand fluteddisk, oocysts ovoidto biconical, in lateralchains. Anthorhynchus
. .
Anthorhynchus longisporaOrmieres & Baudoin
Anthorhynchussophiae(Schneider)
4b. Epimerite a large flattened centrally indented papilla with crenulate border, lost early. Protomerite
with numerous vertical laminations, broadening to an umbrella in the mature sporont, each costule
curved to form a spine pointingbackward; oocystsbiconical orovoid, unitedas a stringofbeads
. .
Sciadiophora
Sciadiophora caudata (Rossler)
Sciadiophorafissidens (Rossler)
Sciadiophoragagrellula Devdhar&Amoji
Sciadiophorageronowitschi(Johansen)
Sciadiophoraphalangii(Leger)
Sciadiophora claviformisOrmieres & Baudoin
5a. Epimerite a conical knob, dentated at the base with a series (about 20) ofvertical lamelle. Oocysts
cylindrical with pointed ends, a tuftofspines ateach pole Contospora opalniaeDevdhar& Amoji
5b. Epimerite simple, globular, without ornamentation 6
6a. Oocystsbarrel-shaped, asymmetrical, withoutterminaltufts, with twoequatorial (lateral)thickenings
on longitudinalcordons Doliospora
Doliospora repelini(Leger)
Doliospora troguli(Geus)
6b. Oocystsbiconical and symmetrical ... 7
7a. Oocystswith 8 to 10 slenderspines at each pole and released in chains of2 to 3 or more from the
gametocyst Echinoocystaphalangii(Amoji & Devdhar)
7b. Oocystswith slenderspines on poles and sides; released unattached from thegametocyst
Cosmetophilus vononesCokendolpher
Order Eugregarinorida Superfamily Stenophoricae
Suborder Septatorina Family Actinocephalidae
Superfamily Gregarinicae Subfamily Actinocephalinae
Family Hirmocystidae
ActinocephalusSteinisarelativelylargegenus
Arachnocystis Levine is restricted to Oribatei with about 40 described species in insects and
mite and opilion hosts. Four species are known, one in an opilion (Levine 1985). Actinocephalus
ofwhichone(thetypespecies)occursinopilions megabuni Ormieres & Baudoin (1973) was dis-
(Levine 1979, \9%5).Arachnocystisarachnoidea covered in the intestine ofa Phalangiidae, Me-
(Devdhar & Gourishankar 1971) was originally gabunusdiadema (Fabricius). Itwascollectedin
described in the genus Sycia Leger (Family Le- Besse, France.
cudinidae).Levine(1979)transferredthespecies Anthorhynchus Labbe are found in a termite
to his new genus Arachnocystis, where it was (Kalavati & Narasimhamurti 1978) and opilion
designated the type species. This species was hosts. Two species are described from harvest-
found in the intestinal ceca of an Assamidae, men. The type species, by monotypy, isAnthor-
Oppalnia sp. (reported as Opalnia sp., see Cok- hynchussophiae Schneider; an opilion parasite.
endolpher 1991), from Someshwar near Dhar- Anthorhynchus longispora Ormieres & Bau-
war, Karnataka State, India (Devdhar 1962). doin (1973) was described from the guts oftwo
.
126 THEJOURNALOFARACHNOLOGY
familiesofharvestmen: Sclerosomatidae [Leiob- describedopilionparasites,thisspeciesisknown
unum (reported as Liobunum) rotundum] and (and illustrated) by all life stages.
Phalangiidae [Mitopus morio, Opilioparietinus, Sciadiophora geronowitschi (Johansen 1894)
Platybunus bucephalus (C. L. Koch)]. All har- was described in the genus Actinocephalus and
vestmen were collected near Besse-en-Chan- was transferred to its present combination by
desse, France. Labbe (1899). This protozoa was discovered in
Anthorhynchus sophiae (Schneider 1887) was theintestinesofaPhalangiidae,Phalangiumop-
originally described in the genus Anthocephalus ilio, from Russia (formerly Russian Soviet Fed-
Schneider. Because that generic name is preoc- erative Socialist Republic, USSR).
cupied, Labbe (1899) provided the replacement Sciadiophoraphalangii (Leger 1897) was first
name and transferred the species to its present described in combination with Lycosella Leger.
combination. The originalcollectionofthispar- It was the type and only species in the genus.
asite was from the intestine of a Phalangiidae, BecauseLycosellawaspreoccupied,Labbe(1899)
Phalangium opilio, capturedin Poitiers, France. proposed the new name Sciadiophora, with S.
This species is also reported by Pfeifer (1956, phalangiihtm%thetype-species.Thisspecieshas
fromGermany)andOrmieres&Baudoin(1973, beenrecorded,redescribedandillustratedbynu-
from France) from the Sclerosomatidae [Leiob- merous authors (Minchin 1903; Wellmer 1911;
unumblackwalli(reportedasL. hassiaeMuller), Ellis 1913;WatsonKamm 1922;Stipperger 1928;
Leiobunum {=Liobunum) rotundum] and the Pfeifer 1956; Silhavy 1961; Geus 1969; Kudo
Phalangiidae [Lacinius ephippiatus, Mitopus 1971; Ormieres & Baudoin 1973). Two families
morio, Oligolophus tridens, Phalangium opilio, ofharvestmen have been reported as hosts (all
Rilaena (reported as Platybunus) triangularis European): Sclerosomatidae [Leiobunum rotun-
(Herbst)]. dum] and Phalangiidae [Lacinius dentiger, Mi-
Sciadiophora Labbe are restricted to opilion topusmorio(reportedasOpiliogrossipes), Opilio
hosts. There are five described species. Devdhar parietinus, Phalangium sp., Phalangium opilio
& Amoji (1978a) provided a table ofcharacters (reported asPhalangium cornutum), Platybunus
which is useful in making identifications. bucephalus, Platybunuspinetorum (C. L. Koch),
Sciadiophoracaudata(Rossler 1882)wasorig- Rileana Platybunus) triangularis]. The origi-
inallydescribedinthegenusStylorhynchusStein. nal collection ofthis species was recorded from
Becausethatnamewaspreoccupied, Ellis(1912) two hosts, Phalangium crassum Dufour and P.
provided the new generic name Stylocephalus cornutum from Vallee de la Loire (where it was
Watson Kamm (1922) transferred the species to rFarraen)cea,nAdsPnrootvedenacbeove(,whtehreelaitttewrasspeccioesmmisonno)w,
itspresentcombinationwithSciadiophora, This known as P. opilio but the identification ofthe
species was originally found in the intestines of
former species is uncertain (Roewer 1923), Re-
Phalangiidae [Mitopusmorio, Odiellus(reported cords ofthis parasite are from France, Austria,
as Odius) spinosus (Bose), Phalangium opilio, Germany and Czechoslovakia.
Phalangiidaegen. sp.] from Germany. Ormieres SciadiophoraclaviformisOrmieres&Baudoin
& Baudoin (1973) reported collections from the (1973) was found in the intestine of a Phalan-
same three hosts from Besse and Tamarissiere, giidae, Mitopus sp. (based on the collection lo-
France. cality the species is probably M. morio). The
Sciadiophorafissidens(Rossler 1882) was first collection locality was Vallee de Chaudefour,
described in the genus Actinocephalus and was France.
later transferred by Labbe (1899) to Sciadi-
ophora. This species was found in intestines of
Subfamily Acanthosporinae
Phalangiidae [Lophopilio (reported as Odiellus)
palpinalis, Phalangiumopilio, Phalangiidaegen. ContosporaDevdhar&Amojiareknownonly
sp.] from Germany. from opilions. The single species, Contospora
Sciadiophora gagrellula Devdhar & Amoji opalniae Devdhar & Amoji (1978a), was de-
(1978b) was described from a Sclerosomatidae, scribed from the midgut and cecum of an As-
Gagrellula saddlana (Roewer), which were col- samidae, Oppalnia sp. (reported as Opalnia sp.,
lected in Dharwarand Kumta, Karnataka State, see Cokendolpher 1991). It is known from So-
India (Devdhar 1962), This gregarine is found meshwarandKalghatgi, DharwarDistrict, India
in the intestine and intestinal ceca. Unlike most (Devdhar 1962).
COKENDOLPHER-=PATHOGENSAND PARASITES OFOPILIONES 127
Cosmetophilus Cokendolpher is a monotypic &Baer(1935)wasdescribedfromtheshrewCro-
genus restricted to an opilion host. It is the only cidura russula Herm. (Insectivora: Soricidae). A
genus of gregarines positively identified from flea, Ctenophtalmus arvernus (Hystrichopsylli-
harvestmen in the New World. Cosmetophilus dae), isknowntobeanintermediatehostofthis
vononesCokendolpher(1991)wasdescribedfrom cestode. Gabrion (1977) reported finding cysti-
the intestine and intestinal ceca ofthe cosmetid cercoides in a Phalangiidae, Phalangium opilio,
harvestman Vonones sayi (Simon) from Texas, collectedduringearlyJuly. Theharvestmanwas
USA. Other samples presumably ofthis species foundinashrewnest(previouslynamedspecies).
were recorded from the same host collected in Shrews in the general area ofthe nest revealed
Tennessee, USA. Unlike mostdescribedopilion proglotids of this cestode as well. It has been
gregarines,thisspeciesisknown(andillustrated) proposed that P. opilio will serve as the inter-
by all life stages. mediate host when fleas are absent.
DoliosporaOrmieres&Baudoin(1969)arere-
strictedto opilionhosts. There are two described Class Trematoda
Order Digenea
species. Doliospora repelini (Leger 1897) was
originally described in the genus Acanthospora Creplin(1846, p. 156) reported findingan un-
Leger (1892). It was designated the type-species identified larval fluke in a Phalangiidae {Phal-
of the new genus, Doliospora, by Ormieres & angium opilio). The fluke was listed as ''Disto-
Baudoin (1969), This species has been reported mum Cystidicola Cr. sp. n.” As no illustration
byLeger(1897)andOrmieres&Baudoin(1973) or description was provided this name must be
from France from the intestines ofthe Sclero- considered a nomen nudum.
somatidae[Leiobunum{=^Liobunum)rotundum]
Family Dicrocoeliidae
andthePhalangiidae[Oligolophustridens, Opilio
parietinus, Megabunusdiadema, Mitopus morio Brachylecithum Strom was originally de-
{=0. grossipes), Phalangium opilio {=P. cornu- scribed as a subgenus of Lypersomum Looss.
turn - type host), Platybunus bucephalus]. Adult flukes ofthis genus are found in the liver
Doliospora troguli (Geus 1969) was originally andgallbladderofbirdsandrarelyinmammals.
described in the genus Acanthospora and trans- Data are available on the life cycles ofsix (in-
ferred to its current combination by Levine complete data for five species) Brachylecithum
(1980). It was found in the intestine of a Tro- spp. (see Carney 1970, 1972). In a typical life-
guloidae, Trogulustricarinatus(Linne),inRaths- cycle the eggs are passed in the feces ofthe de-
berg, Germany. Neither its gametocysts nor its finitive host, a bird or mammal. The eggs are
oocysts are described. eatenbyaterrestrialsnail,theintermediatemol-
Echinoocysta Levine (1984) is composed ofa luscan host, where they develop into miracidia
singlespeciesthatisrestrictedtoanopilionhost. and sporocysts. The cecariae emerge from the
Echinoocystaphalangii(Amoji&Devdhar 1979) snail as a slime ball and are eaten by a second
was originally described in the genus Echinos- intermediate host (usually an arthropod). The
poraAmoji & Devdhar(1979). Because thatge- cecariae encyst in the arthropod hemocoel and
nus was preoccupied, Levine (1984) proposed infect the vertebrate host upon eating the inter-
the new name Echinoocysta and transferred the mediate host. In some arthropod hosts, the me-
speciestoitscurrentcombination.Thisprotozoa tacercariaelodgeinornearthehostbraincausing
isfoundintheintestineandintestinalcecaofan behavioralandmorphologicalchanges(Hohorst
Assamidae, Oppalnia sp. (reported as Opalnia & Graefe 1961; Carney 1969). These changes
sp., see Cokendolpher 1991) from Someshwar, appeartoincreasethechancesofpredationupon
near Dharwad, Karnataka State, India. the arthropod host (Carney 1969).
Brachylecithum sp. cysts and metacercaires
Subkingdom Eumetazoa
were reportedfrom a Phalangiidae, Phalangium
Phylum Platyhelminthes opilio,byGabrion&Ormieres(1973).Thetrem-
Class Cestoda (Cestoidea) atodes were found in the muscles and adipose
Order Cyclophyllidea tissue ofthebody. The infectedharvestmanwas
Subclass Eucestoda
collected in Sete and Montpellier, France. Bra-
Family Hymenolepididae
chylecithum adults are known from Passeri-
Pseudhymenolepis Joyeux & Baer (1935) is formes birds in the south of France. Because
monotypic. Pseudhymenolepis redonica Joyeux Brachlecithum spp. appearto be relatively host-
THEJOURNALOFARACHNOLOGY
128
specific in the arthropod stage ofdevelopment bacteriumXenorhabdusnematophilus(Poinar&
(Carney 1970),therecordinPhalangiumisprob- Thomas) killed the above mentioned arthropod
ably ofan undescribed species. host. This nematode has a wide host range of
insects and arachnids (Poinar 1979; Poinar &
Phylum Nematoda (Nemata) Thomas 1985; Poinar et al. 1985). A thorough
Unidentified Class description and review ofthis nematode and its
Laniatores (Triaenonychidae and/or Synthe- relationship with X. nematophilus are provided
tonychidae) from New Zealand are reported by by Poinar (1979).
Forster(1954)tobeinfestedbyunspecifiednem-
atodes. Dr. V. Tood (in Sankey 1949a) recorded Family Heterorhabditidae
nematodes from Rilaena {=Platybunus) trian- HeterorhabditisPoinaristheonlygenusinthe
gularis. family. It is known by three described species
(Poinar & Welch 1981; Poinar 1990), one of
Class Secementia which is known to infect harvestmen. Keys to
Subclass Rhabditia
theinfectivejuvenilesofthethreespeciesisfound
Order Rhabditida
in Poinar (1990). Adults are identified by elec-
Suborder Rhabditin
trophoreticanalysis ofenzymes (Akhurst 1987),
Superfamily Rhabditoidea DNA fingerprinting and morphology (Poinar et
Family unidentified al. 1987). The mode ofentry into the host and
Pfeifer(1956)reported“rhabditid”nematodes general life cycle follow that listed under Stei-
from Phalangiidae {Lacinius horridus and Phal- nernema,exceptthatHeterorhabditishaveahet-
angiumopilio)thatwerecapturedinBerlin,Ger- erogenic life cycle. Maturing females can either
many. behermaphroditicoramphimictic.Thefirsther-
maphroditic generation is usually followed by
Family Steinemematidae
oneormoreamphimicticgenerationsina single
Steinernema Travassos is comprised of nine cadaver. Juveniles ofHeterorhabditis can enter
distinct species (Poinar 1990). Until recently, host via natural openings, or in smaller, more
most species were referred to Neoaplectana fragile host by breakingthe cuticle with a dorsal
Steiner. Others referred to Neoaplectana are ei- (and sometimes ventral) hook.
thersynonyms,misidentifiedorinsufficientlyde- Heterorhabditis bacteriophora Poinar 1975, is
scribed (Poinar & Welch 1981). Keys and other a well-known insect parasite. Considerable lit-
descriptive data needed for identification ofthe erature on this speciesislistedundera synonym
various species can be located in Poinar (1990). Heterorhabditis heliothidis (Khan et al. 1976);
Onlyonespeciesisknownfromaphalangidhost. which was originally described in combination
Allspeciesthusfardiscoveredcarryasinglespe- with the new genus Chromonema (Khan et al.
cies of symbiotic bacterium in the alimentary 1976). Poinar & Thomas (1985) demonstrated
tract of the third-stage juvenile. The infective this nematode could infect and successfully re-
stagenematodesoccuron soilandhavetheabil- produce in the Phalangiidae Phalangium opilio
itytolocateandenterarthropodhosts. Toreach (reportedasP. sp.). Its symbioticbacterium Xe-
thehemolymphofthehost,thenematodesenter norhabdus luminescens killed the above men-
viaanaturalopeningandthenpenetratethrough tionedhost.Thisnematodehasawidehostrange
the gut or tracheal walls. Once inside the host, ofinsectsand arachnids (Poinar 1979; Poinar&
the nematode releases its associated bacterium Thomas 1985; Poinar et al. 1985). A thorough
which kills the host within 48 hours. The nem- review ofthis nematode is provided by Poinar
atodesmature into males and femalesinsidethe (1979).
arthropod and the females release eggs within
Class Adenophorea
the cadaver (Poinar 1983).
Steinernema carpocapsae (Weiser 1955) was Subclass Enoplia
Order Mermithida
originally described in combination with Neo-
aplectana from codling moth larvae collected in Superfamily Mermithoidea
Czechoslovakia. Poinar & Thomas (1985) dem- Family Mermithidae
onstrated this nematode could infect and suc- All known mermithid records from harvest-
cessfully reproduce in a Phalangiidae, Phalan- men are based on juvenile nematodes. Conse-
gium opilio (reported as P. sp.). Its symbiotic quently, none can be accurately assigned to a
COKENDOLPHER~=PATHOGENSAND PARASITES OFOPILIONES 129
genus (see below under Agamomermis). Re- (1819), new combination, was originally de-
searchers fortunate enough to obtain adult ma- scribed in combination with Filaria. Steiner
terial shouldconsultthekeyprovidedbyPoinar (1922) transferred the species to Mermis. The
(1977). original specimens were from the abdomens of
Matthiesen (1974) reportedthediscoveryofa Phalangiidae, Phalangium opilioand Opilio(re-
Gonyleptidae {Gonyleptesfragilis Mello-Leitao) portedasPhaiangiicornutiand Opilionis). Dies-
which was infested by a internal parasite. Pre- ing (1851) listed the species as ""Gordius trunc-
liminaryobservationsthroughtheharvestman’s tulus Diesing,” but it is unclear if he had
body (the parasite was apparently not dissected additional material
fromthehost)suggestedtheparasitetobeeither AgamermisincertawasreportedbyStipperger
a Nematomorpha or mermithid. Because there (1928) from Mitopus morio collected in Tirol,
are no other recorded cases of the former at- Austria. Stipperger (1928, p. 60) stated that he
tacking Opiliones, I assume the parasite was a hadsentthespecimentoDr. G. Steinerforiden-
juvenile mermithid. tification and that he had received an identifi-
Unknown species were reported by Poinar cation as ""Agamermis incerta n. sp.” Pfeifer
(1985) from a Sclerosomatidae [Togwoteeus(re- (1956)andPoinar(1985)referredto thisspecies
ported as Homolophus) biceps (Thorell) from asAgamermisincerta(Steiner), indicatingthatit
western Canada], a Cosmetidae [Paecilaemana had been described in some othergenus. I have
quadripuncta Goodnight & Goodnight from beenunabletolocatethedescriptionofthisspe-
CostaRica]andaProtolophidae[Protolophussp. cies (in combination with Agamermis Cobb,
from the southwesternUSA]. Pfeifer(1956)also Steiner&Christie;HexamermisSteiner,orMer-
reported an unknown species from a Phalangi- misDujardin)inZoologicalRecord(1918-1940)
idae, Phalangium opilio, from Berlin, Germany. and presume it is a nomen nudum. Apparently,
Tsurusaki (1986) found unidentified mermi- Poinar(1985)alsowasunableto locateSteiner’s
thids in two species ofSclerosomatidae {Leiob- descriptionofincerta(incombinationwithAga-
unum globosum, Leiobunum manubriatum) in mermis or otherwise) from a spider, as this spe-
Japan. cies of mermithid does not occur in his table
Mitov reported (pers. commun.) that he had except associated with Stipperger’s 1928 paper.
discovered larval mermithids in preserved ma- Hexamermissp., incertaesedis,juvenileswere
terial ofthe followingharvestmen from Vitosha reported (Unzicker & Rotramel 1970) from an
Mountain and West Rodopy, Bulgaria: Nemas- immature Phalangiidae harvestman {Opilio sp.-
tomatidae {Paranemastoma radewi), Phalangi- only species in region is O. parietinus) from Il-
idae {Lacinius ephippiatus, L. horridus, L. den- linois, USA. Because of the uncertainty of the
tiger, Lophopilio palpinalis, Mitopus morio, identification,thisspeciesisbestretainedasAga-
Phalangium opilio, Zacheus crista). momermis sp.
Agamomermis Stiles is a collective group Mermis sp. was reported by Kastner (1928).
erected to receive species which were described HestatedJuliusRiihmofNembergsawa“Mer-
from larvae (which lack meaningful taxonomic mis” emerge from a “Phalangiidaen.” This rec-
characters) [see Poinar & Welch (1981)]. When ord was later cited as Phalangiidae, Opilio sp.,
diagnosing Agamomermis, Stiles (1903) stated byPoinar(1985). TheonlypaperbyRiihmcited
byKastnerwaspublishedin 1926andcontained
thegroupwasartificialandthereforeshouldhave
no mention ofa Mermis. Apparently, there has
no type species. All ofthe mermithids thus far been some miscommunication regarding this
recordedfromharvestmen areconsidered incer- record.Probably,Ruhmverballycommunicated
tae sedis and therefore those species that were this observation to Kastnerand used “Mermis”
originallydescribed from harvestmen should be
as a general term for a mermithid nematode.
transferred to Agamomermis. This action was Furthermore, because the record is of a post-
indicatedbutnot formally performed by Poinar
parasiticjuvenile, the record is correctly attrib-
(1985). uted to Agamomermis sp.
Agamomermis phaiangii (Haldeman 1851),
new combination, was originally described in Phylum Nematomorpha
combination with Filaria Mueller from a Phal- Hairworms are free-living as adults and par-
angiidae, Phalangium opilio (reported as P. cor- asitic as juveniles in insects, spiders and crus-
nutum). Agamomermis truncatula Rudolphi taceans. Some early records ofmermithid nem-
