pathogens and parasites of opiliones (arthropoda… · 2004-09-27 · pathogens and parasites of...

1993 . The Journal of Arachnology 21 :120—14 6

PATHOGENS AND PARASITES OF OPILIONES(ARTHROPODA: ARACHNIDA)

James C. Cokendolpher 1 : Adjunct Professor, Department of Biology, MidwesternState University, Wichita Falls, Texas 76308 USA .

ABSTRACT. This is the first paper to review the literature records on all pathogens and parasites of Opilione son a global level . These organisms (bacteria, fungi, protozoans, cestodes, trematodes, nematodes, arthropods )are listed in phylogenetic order along with available information on hosts, collection localities, life history, andtaxonomic history . The opilion hosts are also listed (by their currently accepted names) along with the namesof their known pathogens and parasites . Diagnostic characters and some taxonomic keys are provided for taxawhich are relatively well know . Citations to other available keys are provided . Many new host and distributionrecords are provided.

Two fungi [Engyodontium aranearum (Cavara), Torrubiella pulvinata Mains] are removed from the list ofpathogens of opilions and it is suggested that the original hosts were misidentified spiders.

Two new combinations are recorded in the Mermithidae : Agamomermis phalangii (Haldeman 1851), Aga-momermis truncatula (Rudolphi 1819) . Agamermis incerta Steiner in Stipperger 1928 is regarded as a nomennudum .

The type locality of the mite Leptus lomani (Oudemans 1903b) is restricted to Corral (39°53'S, 73°25'W) ,Valdivia, Chile.

Unlike many arachnids, Opiliones or harvest- ternal pathogens and parasites (see Holmbergmen lack a pumping stomach and therefore theychew their food and often consume oocysts andspores . Examination of their feces reveals a va-riety of chitinous fragments from their arthropodprey as well as plant pieces . Some saprophyticfungi and yeast spores can be observed as wellas gametocytes of internal parasites. The fre-quent grooming of the legs by the harvestmenmay also lead to the ingestion of oocysts andspores. While ingestion is the common entrancepathway for some opilion pathogens, fungi infec ttheir host through penetration of the cuticle . Al -though gregarines and mites are frequently en -countered when observing harvestmen, relative-ly few researchers have documented theiroccurrences .

Harvestmen are unique among arthropods bypossessing bilateral exocrine glands which openonto the dorsal surface of the cephalothorax nearthe base of the second pair of legs. These glandsproduce a variety of volatile secretions (Ekpa etal . 1984, 1985) that have been generally consid-ered to be defensive in nature. The glands havealso been proposed to function in a variety ofother behaviors including protection from ex-

'Home address: 2007 29th Street, Lubbock, Texas79411 USA .

1986, and citations therein) . To date, only de-fense against predators and harvestman aggre-gation formation have been demonstrated .

While working with a South America har-vestman, Estable et al. (1955) discovered tha tthe exocrine gland secretion was a remarkablyeffective antibiotic, in vitro, against 18 genera ofbacteria (Gram positive and negative) and pro -tozoa. Their work revealed that the secretion wasalso active when given orally to mice infectedwith intestinal parasites. The substance was tol-erated perfectly by the mice but destroyed giar-dias, trichomonas and hexamites . The compo-nents of the secretion were later determined t obe a composed of a variety of quinones (Fiese r& Ardao 1956) .

The major components of the exocrine secre-tions of harvestmen differ between the two sub -orders, Laniatores and Cyphopalpatores . Minorcomponents and ratios of components differamong congeneric species (Ekpa etal . 1985). Thefew chemical analyses thus far reported (see Ekp zet al . 1984, and citations therein) from Laniatores reveal a variety of alkylated benzoqui inones, phenols, N,N-dimethyl-,6-phenylethy lamine and bornyl esters. Only the Palpatoresection of the suborder Cyphopalpatores has be echemically investigated. Those analyses reve:

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COKENDOLPHER—PATHOGENS AND PARASITES OF OPILIONES

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members of this group secrete short-chained acy- topic of phoresy arises . Phoresy is not parasitismclic ketones, alcohols and naphthoquinones (see but rather a form of symbiotic relationship inEkpa et al. 1985, and citations therein) .

which the smaller organis mEven though harvestmen are abundant in warm

moist situations, few records are available of fun-gi infecting these animals . Because the majorcomponents of harvestman exocrine secretion sare members of chemical classes known to b efungicides (see Torgeson 1969; Cole et al . 1975),it is likely these secretions are used to protec tharvestmen from infection. The use of these se-cretions in defense and grooming needs furthe rstudy .

While there are several world-wide taxonomi crevisions of harvestmen, no similar treatment fo rtheir parasites has been undertaken . This is inpart due to the incorrect view that harvestme nare not of economic importance . Mounting ev-idence demonstrates that harvestmen are bene-ficial and that they consume considerable quan-tities of pest insects and mites . Because of thisbeneficial status, no one has investigated para-sites for controlling opilions. Experiments in-volving insect pathogens on harvestmen revea lopilions are susceptible . Like conventional in-secticides, insect pathogens and parasites coul dhave a severe impact on the beneficial harvest -men.

Many of the records of parasites from har-vestmen are incomplete . In some cases the host ,but not the parasite, is identified to species . Inother cases, the parasite but not the host is iden-tified to species. The purpose of this contributionis to bring together the limited information onthis topic so that a foundation can be built forfuture research .

Because of the lack of good characters in som egroups (i . e ., Microsporida and juveniles of Mer-mithidae) collective groups have been named .Such groups or genera often include species whichprobably are not related. This group name is usedsimply for "taxonomic convenience" and in-cludes species not readily placed in known genera(possibly because a particular life stage is un-known) and species incertae sedis . Some taxo-nomically convenient groups also occur at higherlevels in fungi . In fungi, the sexual stage (teleo-morph of ascomycetes and basidiomycetes) an dtheir asexual stages (anamorphs or conidial stages)are sometimes placed in separate genera andclasses . In some cases, two or more ascomycetesmay be identified as having the same form spe-cies for an anamorph .

When one discusses parasites of opilions, the

associates with theharvestman in order to obtain transportation.Phoresy as well as passive transport of fungaland plant spores will not be examined here .

Superkingdom ProkaryotaeKingdom Monera

Division Gracilicute sFamily Enterobacteriaceae

Xenorhabdus Thomas & Poinar contains fivedescribed species and other undescribed specie s(Akhurst & Boemare 1990) . They typically in -habit nematodes and their host arthropods (in -sects and arachnids) . See under Nematod a(Rhabditoidea) for further details on this rela-tionship . Pertinent taxonomic papers are citedwith a review of the taxonomic problems in Ak -hurst & Boemare (1990) .

Xenorhabdus luminescens Thomas & Poinar(1979) is introduced into the arthropod host bya Heterorhabditidae [Heterorhabditis bacterio-phoraPoinar] . Poinar & Thomas (1985) dem-onstrated this bacterium could kill a Phalangi-idae (Phalangium opilio Linn., reported as P . sp . )if introduced by the correct nematode.

Xenorhabdus nematophilus (Poinar & Thomas1965) was originally described in combinationwith Achromobacter Bergey, Breed & Murray .This bacterium is introduced into the arthropo dhost by a Steinernematidae [Steinernema car-pocapsae (Weiser)] . Poinar & Thomas (1985)demonstrated that this bacterium could kill aPhalangiidae, Phalangium opilio (reported as P .sp.), if introduced by the proper nematode .

Superkingdom EukaryotaeKingdom Fungi

Division Eumycota

At least one species of fungus successfully kill sa Gonyleptidae (see under Torrubiella gonylep-ticida and unidentified fungi) . Gonyleptoidea areknown to have phenols which are antagonisti cto fungal growth in their exocrine secretions . Ei-ther T. gonylepticida and another unidentifiedfungus from Panama are not retarded by phenols,or the hosts were unable to produce phenols i nsufficient quantity . The extent of phenol pro-duction in various gonyleptid genera and its usein controlling fungi have not been investigated .Likewise, the effects of age and health of the har-

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THE JOURNAL OF ARACHNOLOGY

vestman on phenol production have not beenexamined .

Unidentified Fung i

Griffiths (1978) illustrated a harvestman [no tidentified, but almost certainly Nelima paessler i(Roewer)] covered by mycelia of soil microfungi.The fungi are reported not to be pathogenic, bu tsimply use the harvestman corpse as a substrate .

Mora (1987, fig . 8) reported mortality in adultmales of a Gonyleptidae (Zygopachylus albom-arginis Chamberlin) by an unidentified funguson Barro Colorado Island, Panama . Males of thisnest-building harvestman eat all fungus appear-ing in the nest, thus preventing the proliferationof mycelia . Mora (1987) suggested the males in-gested the fungi (spores) which eventually kille dthem. This is probably incorrect because nearlyall other fungal pathogens of invertebrates infec ttheir host through the cuticle (Samson et al. 1988) .Ten fatalities were observed from 199 nest -guarding males examined by Mora .

Spicaria longipes; which is now recognized asPaecilomyces farinosus (Holm ex S. F. Gray)(Brown & Smith 1957). Petch recorded T. gon-ylepticida and the conidial stage from variousspiders from Trinidad . Koval' (1974) listed theconidial stage from spiders collected on Mag-nolia Linn& leaves in Russia (formerly RussianSoviet Federative Socialist Republic, USSR) . Inthe key to Torrubiella spp . by Koval', two va-rieties of T . gonylepticida are differentiated onthe basis of perithecia and ascus lengths . How-ever, the two taxa should be attributed to anotherspecies : the third taxa in the key should be T .arachnophila var. pleiopus Mains and the fourthshould be T. arachnophila var. pulchra Mains.

Torrubiella pulvinata Mains (1949) was de-scribed from "Opilionoidea" collected on Oahu ,Hawaii. Paecilomyces (reported as Spicaria) pul-vinata (Mains 1949) was the name given to theconidial stage . Samson (1974) listed S . pulvinat aas a synonym of P. farinosus, thus regarding theanamorph for both T. gonylepticida and T. pul-vinata to be the same species. Mains (1949, p .303) stated "The hosts of this collection are soseverely parasitized that accurate determinationis difficult . They appear to be arachnids belong-ing to the Opilionoidea." Because opilions ap-pear to be absent from the Hawaiian Islands (F .G. Howarth pers. commun.), the host is morelikely a long-legged, pholcid spider . The setae -spines on the legs illustrated by Mains (1949, fig .1 A) are long and unlike those on harvestmen .They are similar to those found on spiders. Thereare five adventive cosmopolitan Pholcidae (Ara -neae) established in the islands that could b econfused as opilions by non-specialists . The op-ilion host records are considered here to be in-correct .

Subdivision DeuteromycotinaClass Hyphomycete s

The Hyphomycetes is an artificial class rep-resenting the asexual states of Ascomycetes andBasidiomycetes, or fungi for which sexual state sare unknown . Orders and families do not exist .in current classifications of these fungi .

Hymenostilbe Petch is comprised of seven de -scribed species . Species are known to infect avariety of insect hosts, spiders (Mains 1950 ; Evan s& Samson 1987) and harvestmen . Mains (1950 °stated members of this genus are the conidia l(anamorph) state of Cordyceps spp . ; where&

Subdivision Ascomycotin aClass Pyrenomycete sOrder Clavicipitale sFamily Clavicitaceae

Torrubiella Boudier is a genus with primaryhost affinities for spiders (Araneae), althoug hseveral species are also known from insects, es-pecially Coccidae (Kobayasi & Shimizu 1982 ;Humber & Rombach 1987) . Two species hav ebeen reported from harvestmen, but only onereport appears to be valid .

Torrubiella gonylepticida (Moller 1901) wasoriginally described in combination with Cor-dyceps Fries . Petch (1937) transferred the specie sto its present combination and redescribed thespecies . Moller (1901), when describing the host ,referred to it as a spider ('Die Spinnen', not 'We-berknechte'). Subsequent authors (Petch 1937 ;Koval 1974 ; Kobayasi & Shimizu 1982) hav econtinued to list the only host as a spider . For-tunately, the specific name refers to the true typ ehost, a Gonyleptidae harvestman . Moller (1901 ,taf. 6, fig . 89) clearly illustrated the gonyleptidhost, but not in sufficient detail to determine towhich genus it belongs. Kobayasi & Shimizu(1982) reprinted Miller's illustration and statedthe type locality was Brazil .

Petch (1937) described the conidial stage as


12 3

Evans & Samson (1987) reported the teleomorph

Subdivision Zygomycotinaconnection remains unproven. Specific identifi-

Class Zygomycete scations are best made by consulting the diagnoses

Order Entomophthoralesprovided by Mains (1950) . Hymenostilbe ver-

Family Entomophthoracea erucosa Mains (1950) was originally described Pandora Humber (1989) is comprised of 1 6from spiders collected in Maine, USA . Other re- species of obligately pathogenic fungi . Hosts in-cords are from spiders in England and a "Phal- dude members of insects and arachnids . A singl eangiidae" in England (Leatherdale 1970) .

species is recorded from opilions . Pandora phal-Engyodontium de Hoog is comprised of seven angicida (Lagerheim 1898) was originally de-

species, two of which are reported to infect spi- scribed from Phalangiidae collected in Swedenders and one on "opilionids" . A key to the spe- as a species ofEmpusa Cohn (Entomophthora) .cies is provided by Gams et al . 1984 . Engyo- Batko (1966) transferred the species to Zoo-dontium aranearum (Cavara) was originally phthora Batko 1964, and placed it in the sub -described in the genus Sporotrichum Link exFries 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) recorded this fun-onymy are provided by Gams et al . (1984) . The Sus from a Phalangiidae, Phalangium opilio, inteleomorph state is unknown, but other members England .of the genus have a Torrubiella teleomorph . Those Entomophaga Batko includes 10 describedsame authors reported hosts as a fly, spiders and species (Humber 1989) . All are obligate patho-opilions . The specimen in their photograph (fig . gens of insects and arachnids . A single species is3), as well as those of Samson et al . (1988, pl . recorded from opilions. A key for identification68a,b), superficially resembles opilions, but judg- of members of this genus is provided by Kelle ring from the dense placement and morphology (1987) . Comparisons to original descriptionsof the setae on the host legs (figs . 3, 68b) they (species and citations are listed in Humber 1989 )are not harvestmen . They are more likely pholcid are required for positive identifications . Ento-spiders (Araneae : Pholcidae) . The opilion host mophaga batkoi (Balazy 1978) was originally de -record for this species of fungus is considered scribed in the genus Entomophthora Fresenius.herein incorrect .

Later, Remaudiere & Keller (1980) transferre dNomuraea Maublanc is composed of three de- the species to Conidiobolus Brefeld (Family An-

scribed species (Ignoffo et al. 1989 ; Greenstone cylistaceae), but the current combination wit het al . 1988). Nomuraea rileyi (Farlow) Samson Entomophaga was made by Keller (1987) . Bat-is a well-known pathogen of insects . Nomuraea azy (1978) described this fungus from harvest -atypicola (Yasudo) Samson is reported to infect men collected near Poznan, Poland . Phalangi-spiders, harvestmen and insects . Nomuraea ane- idae [Oligolophus tridens (C. L. Koch)] and rarelymonoides Hocking was originally isolated from Sclerosomatidae (Leiobunum rotundum Latreillesoil and, in high doses in the laboratory, can and Leiobunum blackwalli Meade) were infected .cause mortality in insects .

An epizootic (temporary increase in the inci -Nomuraea atypicola (Yasuda 1915) was orig- dence of infections) was observed during lat e

inally described as a member of the genus Isaria summer .J . Hill ex E. M. Fries . It was found on an Atyp- Keller (1987) reported this species of fungusidae spider in Japan . It was transferred to its was rather common and often caused epizootic spresent combination by Samson (1974) . The te- in open woods, along the borders of forests an dleomorph or sexual state is Cordyceps cylindrica hedges . From late July to the middle of Septem -Petch (1937) . Greenstone et al . (1988) reported ber it was collected from Oligolophus tridens inthe infection of a harvestmen by this fungus un- Switzerland .der laboratory conditions . The infected Sclero -somatidae, Leiobunum vittatum (Say), was col -lected in Missouri, USA . This species of fungus

Kingdom Animaliais commonly found infecting spiders (Green-

Subkingdom Protozoa

stone et al . 1988) and under laboratory condi-

Although seldom reported, Protozoa are corn-tions was also found to be infective to Lepidop- mon parasites of Opiliones . To date, all recordstera larvae (Ignoffo et al . 1989) .

of Protozoan parasites of Opiliones are from USA,

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Europe and India. Their reported absence fromother localities is likely due to lack of study . Whiledissecting gonads for anatomical and chromo-somal studies, I have often observed gregarine sfrom North and Middle America species (es-pecially from Phalangiidae and Cosmetidae) . El-lis (1913, p . 280) reported that he was unable tolocate gregarines in the "alimentary canal of per -haps two hundred Phalangidea" from Michiganand Colorado . His failure to locate parasites mayhave been caused by the time of year he exam-ined the opilions or possibly the taxa he exam-ined do not harbor gregarines (these taxa are un-known, but probably are members of theSclerosomatidae : Leiobuninae as they are th edominate forms in the two mentioned areas) .Only two studies have been published on opilionhematocytes, one of which resulted in the dis-covery of a blood parasite .

Phylum MicrosporaClass Microsporea

Order MicrosporidaCollective Group Microsporidium

Species that cannot be readily placed to genus ,as well as species incertae sedis, are lumped int oMicrosporidium sensu Sprague (1977) . Micro-sporidium weiseri (Silhavy 1960) was originallydescribed in the genus Stempellia Leger & Hesse(Family Thelohaniidae) . Sprague (1977) trans-ferred this species to its present combination withMicrosporidium because the species did not fi tany of the known genera. This parasite was foundin smear-preparations of hemolymph of a Phal-angiidae, Opilio parietinus (De Geer) . The har-vestmen was collected in Trebic, Czechoslova-kia. The plasmodium have 2, 4, 8 and 16 spore sand are found in the hemolymph and hemocyte s(plasmatocytes) of its host.

the feces, and no intermediate host or vector isneeded . Because most species are believed to a t -tach to intestinal epithelial cells, gregarines inopilions probably are not pathogenic .

The gregarine genera and some species know nfrom harvestmen can be identified by the follow -ing taxonomical key. Because some species areinadequately described (some life-stages un-known) identifications to species are difficult .Useful keys or tables of characters are mentione dunder specific genera in the following account.

Class SporozoasidaSubclass Gregarinasin a

Tsurusaki (1986) found gregarines in Sclero-somatidae harvestmen, Leiobunum manubria-turn Karsch and Leiobunum globosum Suzuki ,from numerous localities in Japan . He also pro-vided data on parasitism rates as related to spe-cies, locality and season . His gregarines have notbeen identified .

Hunt (1979) found numerous gregarines in themidgut diverticula of Triaenonychidae harvest -men, Equitius doriae Simon, from southeasternAustralia. His gregarines were never identified .

Mitov reported (pers . commun .) that he haddiscovered gregarines in preserved material o fthe following harvestmen from Vitosha Moun-tain and West Rodopy, Bulgaria: Nemastomatidae [Carinostoma ornatum (Hadzi), Parane-mastoma radewi (Roewer), Pyza bosnica(Roewer)] ; Sclerosomatidae [Leiobunum rume-licum Silhavy] ; Phalangiidae [Lacinius ephippiatus (C . L. Koch), L. horridus (Panzer), L. dentige r(C. L. Koch), Lophopilio palpinalis (Herbst),Mitopus morio (Fabricius), Odiellus lendli (So-rensen), Opilio dinaricus Silhavy, O. ruzicka iSilhavy, O. saxatilis (C. L . Koch), Phalangid mopilio, Zacheus anatolicus (Kulczynski), Z. crist a(Brune)] .

Other new records include unidentified greg-arines from aPhalangiidae, Odiellus pictus Wood,collected in the West Virginia University Forest,Chestnut Ridge, Preston County, WestVirginia ,USA and an unidentified gregarine from a Scle-rosomatidae, Leiobunum politum Weed, collect-ed in Columbus, Ohio, USA. This latter seriesis remarkable as the parasites were only discov-ered after a hundred years of storage .

Phylum Apicomplexa

All known Apicomplexa parasites of opilion sare septate eugregarines and as such have severa lfeatures in common . Both sexual and asexualstages occur (gametogony and sporogony), butmerogony is absent . The mode of infection isingestion of oocysts . The trophozoites attach tothe lining of the gut and divide to form mero-zoites and gamonts . Gametocytes are passed in

Key For Identification Of Gregarines Found In Harvestme n

la . Oocysts without spines or thickening at poles 2lb . Oocysts with spines or thickenings at poles, sometimes at equator and also along edges (Family Actinocephalidae) 5


12 5

Subfamily Acanthosporinae2a. Epimerite simple, spherical ; oocysts biconical, with truncate ends, released unchained by simple de -

hiscense of the gametocyst Family HirmocystidaeArachnocystis arachnoidea (Devdhar & Gourishankar )

2b. Epimerite complex and varied ; oocysts biconical orcylindroconical, united as a string of beads Family Actinocephalidae 3

Subfamily Actinocephalina e3a. Epimerite sessile, with short neck having 8—10 simple digitform processes at apex ; neck persists more

or less in sporont, but digitform processes (tentacles) disappear ; gametocysts dehisce by formation ofhole in wall through which oocysts are extruded in a single thread ; oocysts biconical or lemon -shaped Actinocephalus megabuni Ormieres & Baudoi n

3b. Epimerite without digitform process at apex, gametocysts rupture by simple dehiscence 44a. Epimerite a large, flattened and fluted disk, oocysts ovoid to biconical, in lateral chains . . .Anthorhynchus

Anthorhynchus longispora Ormieres & BaudoinAnthorhynchus sophiae (Schneider)

4b. Epimerite a large flattened centrally indented papilla with crenulate border, lost early . Protomeritewith numerous vertical laminations, broadening to an umbrella in the mature sporont, each costul ecurved to form a spine pointing backward ; oocysts biconical or ovoid, united as a string of beads . . . Sciadiophora

Sciadiophora caudata (ROssler)Sciadiophora fissidens (Rossler)

Sciadiophora gagrellula Devdhar & AmojiSciadiophora geronowitschi (Johansen)

Sciadiophora phalangii (Leger)Sciadiophora claviformis Ormieres & Baudoin

5a. Epimerite a conical knob, dentated at the base with a series (about 20) of vertical lamelle . Oocystscylindrical with pointed ends, a tuft of spines at each pole Contospora opalniae Devdhar & Amoji

5b. Epimerite simple, globular, without ornamentation 66a. Oocysts barrel-shaped, asymmetrical, without terminal tufts, with two equatorial (lateral) thickening s

on longitudinal cordons DoliosporaDoliospora repelini (Leger)

Doliospora troguli (Geus)6b. Oocysts biconical and symmetrical 77a. Oocysts with 8 to 10 slender spines at each pole and released in chains of 2 to 3 or more from th e

gametocyst Echinoocysta phalangii (Amoji & Devdhar)7b. Oocysts with slender spines on poles and sides ; released unattached from the gametocyst Cosmetophilus vonones Cokendolpher

Order EugregarinoridaSuborder Septatorina

Superfamily Gregarinica eFamily Hirmocystidae

Arachnocystis Levine is restricted to Oribateimite and opilion hosts . Four species are known ,of which one (the type species) occurs in opilion s(Levine 1979, 1985). Arachnocystis arachnoidea(Devdhar & Gourishankar 1971) was originallydescribed in the genus Sycia Leger (Family Le-cudinidae) . Levine (1979) transferred the specie sto his new genus Arachnocystis, where it wasdesignated the type species . This species wa sfound in the intestinal ceca of an Assamidae ,Oppalnia sp. (reported as Opalnia sp ., see Cok-endolpher 1991), from Someshwar near Dhar-war, Karnataka State, India (Devdhar 1962).

Superfamily StenophoricaeFamily Actinocephalida e

Subfamily Actinocephalinae

Actinocephalus Stein is a relatively large genuswith about 40 described species in insects an done in an opilion (Levine 1985) . Actinocephalusmegabuni Ormieres & Baudoin (1973) was dis-covered in the intestine of a Phalangiidae, Me-gabunus diadema (Fabricius) . It was collected i nBesse, France.

Anthorhynchus Labbe are found in a termite(Kalavati & Narasimhamurti 1978) and opilionhosts . Two species are described from harvest -men. The type species, by monotypy, is Anthor-hynchus sophiae Schneider; an opilion parasite .

Anthorhynchus longispora Ormieres & Bau-doin (1973) was described from the guts of two

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described opilion parasites, this species is known(and illustrated) by all life stages.

Sciadiophora geronowitschi (Johansen 1894)was described in the genus Actinocephalus andwas transferred to its present combination byLabbe (1899) . This protozoa was discovered inthe intestines of a Phalangiidae, Phalangium op-ilio, from Russia (formerly Russian Soviet Fed -erative Socialist Republic, USSR) .

Sciadiophora phalangii (Leger 1897) was firstdescribed in combination with Lycosella Leger.It was the type and only species in the genus .Because Lycosella was preoccupied, Labbe (1899 )proposed the new name Sciadiophora, with S .phalangii being the type-species . This species ha sbeen recorded, redescribed and illustrated by nu-merous authors (Minchin 1903; Wellmer 1911 ;Ellis 1913; Watson Kamm 1922; Stipperger 1928 ;Pfeifer 1956 ; Silhavy 1961 ; Geus 1969; Kudo1971 ; Ormieres & Baudoin 1973). Two familiesof harvestmen have been reported as hosts (al lEuropean) : Sclerosomatidae [Leiobunum rotun-dum] and Phalangiidae [Lacinius dentiger, Mi-topus morio (reported as Opilio grossipes) ; Opilioparietinus, Phalangium sp., Phalangium opilio(reported as Phalangium cornutum), Platybunusbucephalus, Platybunus pinetorum (C . L. Koch) ,Rileana (=Platybunus) triangularis] . The origi-nal collection of this species was recorded fro mtwo hosts, Phalangium crassum Dufour and P .cornutum from Vallee de la Loire (where it wasrare) and Provence (where it was common) ,France . As noted above, the latter species is nowknown as P. opilio but the identification of theformer species is uncertain (Roewer 1923) . Re -cords of this parasite are from France, Austria ,Germany and Czechoslovakia.

Sciadiophora claviformis Ormieres & Baudoin(1973) was found in the intestine of a Phalan-giidae, Mitopus sp . (based on the collection lo-cality the species is probably M. morio) . Thecollection locality was Vallee de Chaudefour,France .

families of harvestmen : Sclerosomatidae [Leiob-unum (reported as Liobunum) rotundum] andPhalangiidae [Mitopus morio, Opilio parietinus ,Platybunus bucephalus (C . L. Koch)] . All har-vestmen were collected near Besse-en-Chan-desse, France .

Anthorhynchus sophiae (Schneider 1887) wasoriginally described in the genus AnthocephalusSchneider . Because that generic name is preoc-cupied, Labbe (1899) provided the replacementname and transferred the species to its presen tcombination . The original collection of this par-asite was from the intestine of a Phalangiidae ,Phalangium opilio, captured in Poitiers, France .This species is also reported by Pfeifer (1956 ,from Germany) and Ormieres & Baudoin (1973 ,from France) from the Sclerosomatidae [Leiob-unum blackwalli (reported as L. hassiae Muller) ,Leiobunum (=Liobunum) rotundum] and thePhalangiidae [Lacinius ephippiatus, Mitopusmorio, Oligolophus tridens, Phalangium opilio ,Rilaena (reported as Platybunus) triangularis(Herbst)] .

Sciadiophora Labbe are restricted to opilionhosts . There are five described species . Devdhar& Amoji (1978a) provided a table of character swhich is useful in making identifications.

Sciadiophora caudata (Rossler 1882) was orig-inally described in the genus Stylorhynchus Stein .Because that name was preoccupied, Ellis (1912)provided the new generic name Stylocephalus .Watson Kamm (1922) transferred the species toits present combination with Sciadiophora . Thisspecies was originally found in the intestines o fPhalangiidae [Mitopus morio, Odiellus (reportedas Odius) spinosus (Bost), Phalangium opilio,Phalangiidae gen . sp .] from Germany. Ormieres& Baudoin (1973) reported collections from th esame three hosts from Besse and Tamarissiere ,France .

Sciadiophora fissidens (Rossler 1882) was firstdescribed in the genus Actinocephalus and waslater transferred by Labbe (1899) to Sciadi-ophora . This species was found in intestines ofPhalangiidae [Lophopilio (reported as Odiellus)palpinalis, Phalangium opilio, Phalangiidae gen .sp .] from Germany .

Sciadiophora gagrellula Devdhar & Amoj i(1978b) was described from a Sclerosomatidae,Gagrellula saddlana (Roewer), which were col-lected in Dharwar and Kumta, Karnataka State ,India (Devdhar 1962) . This gregarine is foundin the intestine and intestinal ceca . Unlike most

Subfamily Acanthosporinae

Contospora Devdhar & Amoji are known onlyfrom opilions. The single species, Contosporaopalniae Devdhar & Amoji (1978a), was de -scribed from the midgut and cecum of an As-samidae, Oppalnia sp. (reported as Opalnia sp . ,see Cokendolpher 1991) . It is known from So-meshwar and Kalghatgi, Dharwar District, India(Devdhar 1962) .


12 7

Cosmetophilus Cokendolpher is a monotypi cgenus restricted to an opilion host . It is the onlygenus of gregarines positively identified fro mharvestmen in the New World. Cosmetophilusvonones Cokendolpher (1991) was described fromthe intestine and intestinal ceca of the cosmeti dharvestman Vonones sayi (Simon) from Texas,USA. Other samples presumably of this specie swere recorded from the same host collected i nTennessee, USA . Unlike most described opiliongregarines, this species is known (and illustrated )by all life stages .

Doliospora Ormieres & Baudoin (1969) are re -stricted to opilion hosts . There are two describedspecies . Doliospora repelini (Leger 1897) wa soriginally described in the genus AcanthosporaLeger (1892) . It was designated the type-speciesof the new genus, Doliospora, by Ormieres &Baudoin (1969) . This species has been reportedby Leger (1897) and Ormieres & Baudoin (1973 )from France from the intestines of the Sclero-somatidae [Leiobunum (=Liobunum) rotundum]and the Phalangiidae [Oligolophus tridens, Opilioparietinus, ]I/Iegabunus diadema, Mitopus morio(=O. grossipes), Phalangium opilio (=P . cornu-turn - type host), Platybunus bucephalus] .

Doliospora troguli (Geus 1969) was originallydescribed in the genus Acanthospora and trans-ferred to its current combination by Levine(1980) . It was found in the intestine of a Tro-guloidae, Trogulus tricarinatus (Linn), in Raths-berg, Germany . Neither its gametocysts nor itsoocysts are described .

Echinoocysta Levine (1984) is composed of asingle species that is restricted to an opilion host .Echinoocysta phalangii (Amoji & Devdhar 1979)was originally described in the genus Echinos-pora Amoji & Devdhar (1979) . Because that ge-nus was preoccupied, Levine (1984) propose dthe new name Echinoocysta and transferred th especies to its current combination . This protozo ais found in the intestine and intestinal ceca of a nAssamidae, Oppalnia sp . (reported as Opalniasp ., see Cokendolpher 1991) from Someshwar ,near Dharwad, Karnataka State, India .

Subkingdom EumetazoaPhylum Platyhelminthe sClass Cestoda (Cestoidea)

Order Cyclophyllide aSubclass Eucestoda

Family Hymenolepididae

Pseudhymenolepis Joyeux & Baer (1935) i smonotypic . Pseudhymenolepis redonica Joyeux

Class TrematodaOrder Digenea

Creplin (1846, p . 156) reported finding an un-identified larval fluke in a Phalangiidae (Phal-angium opilio) . The fluke was listed as "Disto-mum Cystidicola Cr. sp . n." As no illustrationor description was provided this name must b econsidered a nomen nudum .

Family Dicrocoeliidae

Brachylecithum Strom was originally de -scribed as a subgenus of Lypersomum Looss.Adult flukes of this genus are found in the live rand gall bladder of birds and rarely in mammals .Data are available on the life cycles of six (in-complete data for five species) Brachylecithumspp . (see Carney 1970, 1972) . In a typical life -cycle the eggs are passed in the feces of the de-finitive host, a bird or mammal . The eggs areeaten by a terrestrial snail, the intermediate mol-luscan host, where they develop into miracidi aand sporocysts. The cecariae emerge from th esnail as a slime ball and are eaten by a secondintermediate host (usually an arthropod) . Thececariae encyst in the arthropod hemocoel an dinfect the vertebrate host upon eating the inter -mediate host . In some arthropod hosts, the me-tacercariae lodge in or near the host brain causingbehavioral and morphological changes (Hohors t& Graefe 1961; Carney 1969) . These change sappear to increase the chances of predation upo nthe arthropod host (Carney 1969).

Brachylecithum sp . cysts and metacercaireswere reported from a Phalangiidae, Phalangiumopilio, by Gabrion & Ormieres (1973) . The trem -atodes were found in the muscles and adipos etissue of the body . The infected harvestman wascollected in Sete and Montpellier, France . Bra-chylecithum adults are known from Passeri-formes birds in the south of France . BecauseBrachlecithum spp . appear to be relatively host -

& Baer (1935) was described from the shrew Cro-cidura russula Herm. (Insectivora : Soricidae) . Aflea, Ctenophtalmus arvernus (Hystrichopsylli-dae), is known to be an intermediate host of thi scestode . Gabrion (1977) reported finding cysti-cercoides in a Phalangiidae, Phalangium opilio ,collected during early July . The harvestman wasfound in a shrew nest (previously named species) .Shrews in the general area of the nest revealedproglotids of this cestode as well . It has beenproposed that P . opilio will serve as the inter-mediate host when fleas are absent .

128


specific in the arthropod stage of developmen t(Carney 1970), the record in Phalangium is prob-ably of an undescribed species .

Phylum Nematoda (Nemata)Unidentified Clas s

Laniatores (Triaenonychidae and/or Synthe-tonychidae) from New Zealand are reported byForster (1954) to be infested by unspecified nem -atodes . Dr . V. Tood (in Sankey 1949a) recordednematodes from Rilaena (=Platybunus) trian-gularis .

Class Secernenti aSubclass RhabditiaOrder Rhabditida

Suborder RhabditinSuperfamily Rhabditoide a

Family unidentified

Pfeifer (1956) reported "rhabditid" nematodesfrom Phalangiidae (Lacinius horridus and Phal-angium opilio) that were captured in Berlin, Ger -many.

Family Steinernematidae

Steinernema Travassos is comprised of nin edistinct species (Poinar 1990). Until recently ,most species were referred to Neoaplectan aSteiner. Others referred to Neoaplectana are ei-ther synonyms, misidentified or insufficiently de -scribed (Poinar & Welch 1981) . Keys and othe rdescriptive data needed for identification of th evarious species can be located in Poinar (1990) .Only one species is known from a phalangid host .All species thus far discovered carry a single spe -

bacterium Xenorhabdus nematophilus (Poinar &Thomas) killed the above mentioned arthropo dhost . This nematode has a wide host range ofinsects and arachnids (Poinar 1979 ; Poinar &Thomas 1985 ; Poinar et al. 1985). A thoroughdescription and review of this nematode and it srelationship with X. nematophilus are provide dby Poinar (1979) .

Family Heterorhabditidae

Heterorhabditis Poinar is the only genus in thefamily . It is known by three described specie s(Poinar & Welch 1981 ; Poinar 1990), one ofwhich is known to infect harvestmen . Keys tothe infective juveniles of the three species is foundin Poinar (1990) . Adults are identified by elec-trophoretic analysis of enzymes (Akhurst 1987) ,DNA fingerprinting and morphology (Poinar etal . 1987) . The mode of entry into the host andgeneral life cycle follow that listed under Stei -nernema, except that Heterorhabditis have a het -erogenic life cycle . Maturing females can eithe rbe hermaphroditic or amphimictic. The first her-maphroditic generation is usually followed b yone or more amphimictic generations in a singl ecadaver. Juveniles of Heterorhabditis can ente rhost via natural openings, or in smaller, morefragile host by breaking the cuticle with a dorsa l(and sometimes ventral) hook .

Heterorhabditis bacteriophora Poinar 1975, isa well-known insect parasite . Considerable lit-erature on this species is listed under a synonymHeterorhabditis heliothidis (Khan et al. 1976) ;which was originally described in combinatio nwith the new genus Chromonema (Khan et al.

cies of symbiotic bacterium in the alimentary 1976) . Poinar & Thomas (1985) demonstratedtract of the third-stage juvenile . The infective this nematode could infect and successfully re -stage nematodes occur on soil and have the abil- produce in the Phalangiidae Phalangium opilioity to locate and enter arthropod hosts . To reach (reported as P. sp .) . Its symbiotic bacterium Xe-the hemolymph of the host, the nematodes enter norhabdus luminescens killed the above men-via a natural opening and then penetrate through tioned host . This nematode has a wide host rangethe gut or tracheal walls . Once inside the host, of insects and arachnids (Poinar 1979 ; Poinar &the nematode releases its associated bacterium Thomas 1985 ; Poinar et al. 1985) . A thoroughwhich kills the host within 48 hours . The nem- review of this nematode is provided by Poinaratodes mature into males and females inside the (1979) .arthropod and the females release eggs withi nthe cadaver (Poinar 1983) .

Steinernema carpocapsae (Weiser 1955) wasoriginally described in combination with Neo-aplectana from codling moth larvae collected i nCzechoslovakia. Poinar & Thomas (1985) dem-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

Class AdenophoreaSubclass Enoplia

Order MermithidaSuperfamily Mermithoide a

Family Mermithidae


12 9

genus (see below under Agamomermis) . Re- (1819), new combination, was originally de -searchers fortunate enough to obtain adult ma- scribed in combination with Filaria . Steine rterial should consult the key provided by Poinar (1922) transferred the species to Mermis . The(1977) .

original specimens were from the abdomens o fMatthiesen (1974) reported the discovery of a Phalangiidae, Phalangium opilio and Opilio (re -

Gonyleptidae (Gonyleptes fragilis Mello-Leitao) ported as Phalangii cornuti and Opilionis) . Dies-which was infested by a internal parasite . Pre- ing (1851) listed the species as "Gordius trunc-liminary observations through the harvestman's tulus Diesing," but it is unclear if he hadbody (the parasite was apparently not dissected additional material .from the host) suggested the parasite to be either Agamermis incerta was reported by Stippergera 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 hetacking Opiliones, I assume the parasite was a had sent the specimen to Dr . G. Steiner for iden -juvenile mermithid .

tification and that he had received an identifi -Unknown species were reported by Poinar cation as "Agamermis incerta n. sp." Pfeife r

(1985) from a Sclerosomatidae [Togwoteeus (re- (1956) and Poinar (1985) referred to this specie sported as Homolophus) biceps (Thorell) from as Agamermis incerta (Steiner), indicating that itwestern Canada], a Cosmetidae [Paecilaemana had been described in some other genus . I havequadripuncta Goodnight & Goodnight from been unable to locate the description of this spe -Costa Rica] and a Protolophidae [Protolophus sp. cies (in combination with Agamermis Cobb ,from the southwestern USA] . Pfeifer (1956) also Steiner & Christie ; Hexamermis Steiner, or Mer-reported an unknown species from a Phalangi- mis Dujardin) in Zoological Record (1918–1940)idae, Phalangium opilio, from Berlin, Germany . and presume it is a nomen nudum . Apparently,

Tsurusaki (1986) found unidentified mermi- Poinar (1985) also was unable to locate Steiner' sthids in two species of Sclerosomatidae (Leiob- description of incerta (in combination with Aga-unum globosum, Leiobunum manubriatum) in mermis or otherwise) from a spider, as this spe -Japan .

cies of mermithid does not occur in his tabl eMitov reported (pers . commun .) that he had except associated with Stipperger's 1928 paper .

discovered larval mermithids in preserved ma- Hexamermis sp., incertae sedis, juveniles wereterial of the following harvestmen from Vitosha reported (Unzicker & Rotramel 1970) from a nMountain 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 th e

tiger, Lophopilio palpinalis, Mitopus morio, identification, this species is best retained as Aga -

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 He stated Julius Ram of Nernberg saw a "

from larvae 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 by Poinar (1985) . The only paper by Riihm cited

the group was artificial and therefore should have by Kastner was published in 1926 and containe d

no type species. All of the mermithids thus far no mention of aMermis . Apparently, there has

been some miscommunication regarding thisrecorded from harvestmen are considered inter- record . Probably, Riihm verbally communicate dtae sedis and therefore those species that were this observation to Kastner and used "Mermis"originally described 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 -indicated but not formally performed by Poinar parasitic juvenile, the record is correctly attrib -(1985) .

uted to Agamomermis sp.Agamomermis phalangii (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 of mermithid nem-

130


atodes were incorrectly assigned to two genera(Filaria and Gordius Linne) belonging to thisphylum. Those species are listed herein as Aga-momermis spp . (see this group under the Nem-atoda) .

Phylum ArthropodaClass Insecta

Order DipteraSuborder Cyclorrhapha

The Cyclorrhapha is comprised of many fam-ilies of flies, each having a different life cycle –most are not parasitoides . Without knowing theidentification of the fly, little can be written othe rthan a notice of the single reported occurrence .

Soares (1945) reported the discovery of a fl ypupa inside an adult of a Gonyleptidae (Disco-cyrtus invalidus Piza) . The gonyleptid was col-lected at Porto Cabral, Estado de Sao Paulo, Bra -zil.

Suborder NematoceraFamily Ceratopogonidae

Tsurusaki (pers . commun.) reported findin gadult flies of this family, subfamily Forcipomyi-inae, settled on the leg femora of Nelima nigri-coxa and Gagrellula ferruginea in Japan . Whenhe disturbed the flies they would hover aroun dthe host . He suspected they were sucking bloodfrom the harvestmen.

and unlike other pompilids is not very selectiveas to the prey it takes . Prey items include at least22 species of spiders from seven different fami-lies (Evans & Yoshimoto 1962) . Evans (1948)recorded a female wasp taking a Phalangiidae(Odiellus pictus) in East Hartford, Connecticut ,USA. Because the harvestman was taken awa yfrom the wasp before it dug a nest it is uncertai nif it would use the O . pictus to provision the nest.Evans (pers . commun.) recalled that the was pwas captured while it was dragging the opilionacross the ground but he could not determin ewhether the opilion was used in provisioning thewasp nest . Pompilids often take prey and the nabandon it, sometimes after feeding on it .

Class ArachnidaOrder Acarin a

Suborder Prostigmata

Mites known to be parasitic on harvestme nbelong to the families Thrombidiidae and Ery-thraeidae. Only the larval forms are parasiti c(protelean parasites) while the nymphs and adult sare predaceous on small insects . Because the lar-val and post-larval stages of these two familie sare heteromorphic, systematists have long use ddifferent scientific names for each (Southcott1961) . Only after the larval and post-larval stage sare associated by rearings can any meaningfulclassifications be constructed .

Laniatores (Triaenonychidae and/or Synthe-tonychidae) from New Zealand are reported b yForster (1954) to be often heavily infested bymites . Hunt (1979) found a species of parasiticmite on Triaenonychidae harvestman, Equitiusdoriae Simon, from southeastern Australia . Bur -ton & Burton (1984, pp . 218 and 226) publishe da color photograph of a harvestman with nu-merous parasitic red mites . The harvestman i sclearly Mitopus morio . The mites are probablymembers of the genus Leptus, although this ca nnot be stated for certain . Elliott & Reddell (1987 )reported that many of the Leiobunum townsendioccurring in caves in central Texas carried redchiggers on their legs . The mites are probably no tchiggers but more likely the larvae of Leptus .Eaton (1985) stated in a report on some har-vestmen from a cave in southeastern, New Mex-ico that the "The [harvestmen] spiders all hadone or more small, shiny, bright red, oval bumpson their legs which appeared to be some kind o fparasite ." These parasites are likewise probabl yLeptus sp . and the hosts are almost certainly L .townsendi .

Order Hymenopter aFamily Chalcidae

Laniatores (Triaenonychidae and/or Synthe-tonychidae) from New Zealand are reported b yForster (1954) to be infested by chalcid wasps .No specific identifications were provided .

Family Pompilidae

Anoplius Dufour is a large, diverse group o fwasps which prey almost exclusively upon spi-ders (Evans & Yoshimoto 1962) . The femalewasps sting and paralyze spiders which are in-dividually entombed with a wasp egg. The was pyoung will then devour the spider as it grows .Some of the species permanently paralyze thei rprey while others only paralyze them temporar-ily . Some adult wasps feed upon spiders whil eothers feed upon nectar of flowers . Only a singlespecies has been recorded to prey upon a har-vestman.

Anoplius (Pompilinus) marginatus (Say 1824 )is found over most of temperate North Americaeast of the Rocky Mountains . It is often common


13 1

Other unidentified mites from my collection inal spiracles of a Phalangiidae, Zacheus cristainclude: Cosmetidae (Vonones sayi) from Sam Brulle . The collections were from Lindos, Rho -Houston National Forest, Lake Stubblefield, dos .Walker County, and Lake Kirby, Taylor County, Trombidium Fabricius is a relatively large ge-Texas, USA (mites found on dorsa of abdomens) ; nus of conspicuous mites with about 20 species .Phalangiidae [Zacheus hebraicus (Simon)] from Member species have been observed and re -Beith Shemesh, Israel (mite from tibia I) ; Scle- corded since the first record in about 300 B .C .rosomatidae : Leiobuninae (Leiobunum townsen- by Apollodorus . About half of the described spe-di Weed) from near Cloudcroft, New Mexico, cies are known only by adults . Juveniles areUSA; [Leiobunum ventricosum (Wood)], West known to feed on numerous orders of insects asVirginia University Forest, Chestnut Ridge, well as spiders, a pseudoscorpion and harvest-Monongalia County, West Virginia, USA . ; (Nei- men (Welbourn 1983) .ima paessleri Roewer) from Moose Creek Re- Yokogawa (1940) described and illustrated asearch Station, Idaho, USA ; Sclerosomatidae : Sclerosomatidae (Nelima sp .) parasitized by aGagrellinae (Gagrellopsis nodulifera Sato & Su- mite from Japan . The mite was identified as azuki) from Mt . Daisen, Tottori Pref., Japan (mite "Trombidinium" [sic] . Trombidium hungariumfound on dorsum of abdomen) ; (Trachyrhinus Kobulej (1957) is recorded from a Phalangiidaerectipalpus Cokendolpher) from 2 km W . of (Egaenus convexus Koch) from Matraszentimre ,Cuevitas, Starr County and Buffalo Gap, Taylor Hungary . Both the larva and the nymph of thi sCounty, Texas, USA ; (Prionostemma panama species were described by Kobulej (1957) .Goodnight & Goodnight) from Orillas de Rio

Family ErythraeidaeMata Ahogado el Vallo de Anton, Prov . Code ,Panama (mite was found on the abdomen) ; Scle-

The first record of a harvestman parasite wa srosomatidae : Metopilio Group (Globipes sp .) probably an erythraeid mite . Lister (1678) re -from near Cloudcroft, New Mexico, USA .

ported scarlet-colored "bugs" attached and feed -ing from what is now known to be Phalangiidae

Family Thrombidiidae

Phalangium opilio in England . Sankey (1949b )Known as the velvet mites, adults of this fam- reported mites of this family from numerous spe-

ily are among the largest and most conspicuous cies of harvestmen collected in England . Specif-families of mites. ically, he recorded hosts as : Sclerosomatidae

Allothrombium Berlese is a small genus with [Leiobunum blackwalli, L . rotundum, Nelimaseven described species . Its members are para- silvatica (Simon)] and Phalangiidae [Mitopussitic on harvestmen, spiders, several orders of morio, Oligolophus hansenii (Kraepelin), O . tri-insects and isopods (Welbourn 1983) . Megnin dens, Opilio parietinus, Paroligolophus agresti s(1876) described the larva of a mite reared from (Meade), Phalangium opilio, Rilaena triangu-opilions . He identified the mite as either Trom- laris (–Platybunus triangularis)] . Sankey (1949a)bidium fuliginosum Herman or Trombidium stated that he had records of 10 species of har-gymnopterorum Berlese . Based on the structure vestmen (presumably those listed above) beingof the tarsal claws, Southcott (1961) identified used as carriers by the larvae of Erythraeus phal-Megnin's specimen (which was illustrated) as an angioides (De Geer 1778). This identification isAllothrombium sp .

probably incorrect as this species is not otherwiseAllothrombium chanaanense Feider (1977) was known to feed on harvestmen and there is som e

described from an "Opilionida" from Jerusalem, question regarding the true identity of larval E.Israel . This species of mite is only known from phalangiodes (see Southcott 1961) . Possibly ,the larval forms. Host records also include in- Sankey confused the names phalangii and phal-sects: an Acrididae [Prionsosthenus galericulatus angiodes; both of which were described by De(Stal)] and an unidentified Aphidae from Israel Geer. Martinez Crespo & Morales Soto (1979)(Feider 1977) .

reported that mites of the family ErythraeidaeAllothrombium neapolitum Oudemans (1910a) were parasitic on Opiliones from Mexico .

was described from a Phalangidae (Phalangium There are over 50 species of Charletoniasp .) from Portici, Campania, southern Italy. Oudemans described as larvae and 22 specie sOudemans (1913) redescribed and illustrated this described originally as adults . Species are re -species . Specimens identified from my collection corded from every continent except Antarctic awere found attached to the edges of the abdom- (Southcott 1991) . Larvae of two species are par-

132

THE JOURNAL OF ARACHNOLOG Y

asitic on harvestmen (Kawashima 1961 ; South- host) from the USA and provided some addi-cott 1961, 1965, 1991) . The other species are tional comments on members of the genus .common parasites as juveniles on locusts (Acrid- Southcott (1992) described numerous new spe-idae) and less commonly encountered on jump- cies and provided a key to the taxa from Northing plant lice (Psyllidae : Homoptera), true bugs America and Europe . He also resolved the iden-(Lygaeidae and Miridae : Hemiptera), wasps tity of L. ignotus and found the type species,(Braconidae: Hymenoptera), Lepidoptera, drag- Acarus phalangii, to be an illegitimate name . Ka-onflies (Libellulidae : Odonata), flies (Tabanidae, washima (1958) and Haitlinger (1990) providedDolichopodidae and Bombylidae : Diptera), keys to the parasitic larval forms from Japan an dmantis (Mantidae : Mantodea), walking sticks northern Africa, respectively .(Phasmatidae : Phasmida), katydids (Tettigoni- Leptus spp . have been reported from a varietyidae : Orthoptera), beetles (Curculionidae, Me- of hosts and localities . The mode of attachmentlyridae, Tenebrionidae : Coleoptera), mites (Er- was described in Leptus sp . on two species ofythraeidae : Acarina) and spiders (Theridiidae, Phalangiidae (Mitopus morio, Phalangium opi-Philodromidae : Araneae) . Keys to the species are no) by Abro (1988) . Abro (1991) described un -provided by Southcott (1991).

successful parasitic attachments of larval LeptusCharletonia enghoffi Southcott (1991) is known spp. to the ocular tubercle ofPhalangium . Evans

by four larvae recovered from the dorsum and et al . (1961, fig. 211) illustrated a Phalangiidaefemora of the Phalangiidae, Bunochelis canar- (Mitopus morio) infested with Leptus sp. larvaeiana (Strand) . The species were obtained in Feb- from the British Isles . Welbourn (1983) reportedruary in Teno Barranco de las Cuevas, Tenerife, Leptus spp. from unidentified Opiliones collect-Canary Islands .

ed in Ohio and Arkansas .Charletonia southcotti Kawashima (1961) is Robert G. Holmberg reported (pers . corn -

recorded from a Sclerosomatidae, Metagagrella mun.) that he had found 39 vials of harvestmentenuipes (L . Koch) (reported as Gagrella japonica infested with 78 mites, all of which have bee nRoewer), that was collected at the seashore of identified as Leptus spp. by I . M. Smith (Bio-Kasumigaoka, Fukuoka City, Fukuoka-Prefec- systematic Research Center, Ottawa, Canada) .ture, Kyushu, Japan . This species of mite is only Dr. Holmberg's collections were from : "Tog-known from the single collection on 12 July. woteeus biceps from Canada and the USA, Mi -Thirty-five mites were recovered from 20 opi- topus morio from England, Odiellus pictus fromlions . It is known only by the larval stage, which Canada, Oligolophus tridens from Canada, Par-was redescribed by Southcott (1965).

oligolophus agrestis from Wales, Phalangium op -Leptus Latreille is a large genus and its mem- ilio from Canada and England, and Leiobunum

bers are widespread. Over 60 Leptus spp. have townsendi from U .S .A . "been described from larvae . Many adults have Mullen (1988) reported "opilionids corn-also been described, but only in a few cases have monly serve as the host to Leptus mites. Savorycorrelations been made between larval and post- (1938) recorded Belaustium [sic] (Ritteria) ne-larval forms . Only in a single case is a species morum (Koch) from harvestmen in England .described from larval and all post-larval stages This observation was later cited on several oc-(Welbourn & Jennings 1991) . Many species re- casions in general works about arachnids by Sa-main undescribed . Member species are generally vory and Cloudsley-Thompson . The original ob -parasitic on spiders, scorpions, harvestmen, dip- servations were most likely based on alopods, Collembola, and insects . Many of the misidentification and probably were represen-early reports and even some more recent are sus- tatives of the genus Leptus . Not only is Leptuspect as the true identity of the mites identified widely known as a harvestman parasite, butis uncertain . Southcott (1961, 1991, 1992) re- members of the Balaustiinae are generally con -viewed some of the problems regarding the Eu- sidered not to be parasitic on arthropods (South-ropean mites (phalangii, ignotus, nemorum, coc- cott 1961) .cineus) which had been referred to various genera. Cox et al . (1921) found an immature Ery -Southcott (1989) provided a key to the parasitic thaeus [sic] sp . on a "phalangid" in California ,larval forms that were recognizable (most early USA . This mite is probably a Leptus sp . Mite sdescriptions are inadequate to recognize the spe- from almost every genus of the Erythraeidae havecies) in the New World. Welbourn & Jennings been misidentified as Erythraeus Latreille (see(1991) added a new species (from Lepidoptera Southcott 1961) .


13 3

Leptus phalangii (De Geer 1778) was origi-nally described in combination with AcarusLinne. When erecting the genus Leptus, Latreill e(1796) designated (by monotypy) Acarus phal-angii as the type species . The type specimen swere from a Phalangiidae (Phalangium sp .) col-lected in Sweden . Apparently, none of De Geer'sspecimens were preserved . There has been con-siderable confusion over the identity of this spe-cies . Furthermore, as noted by Southcott (1992 )the specific name is not available under the In-ternational Code of Zoological Nomenclature a sDe Geer did not treat it consistently as a bino-men. Only in a few cases can specimens that hav ebeen previously referred to in the literature a sLeptus phalangii be assigned currently recog-nized names .

Leptus ignotus (Oudemans 1903a) was origi-nally described in combination with Erythraeus .The type locality is Borkum, Holland. Southcott(1991) redescribed the species and limited thespecies diagnosis to specimens which had notbeen recorded from opilion hosts . Therefore al lrecords of this species from opilion host can beassumed to be misidentified and are referred t oin Table 1 as Leptus sp . Evans et al . (1961) re -corded a mite (identified as L. ignotus) parasiticon Opilio parietinus in the British Isles . Otherrecords are also from the Phalangiidae : Mitopusmorio from Tirol, Austria (Stipperger 1928) andBulgaria (Beron 1975) ; Opilio ruzickai from Bul-garia (Beron 1975); and Phalangium opilio, R .triangularis, Lophopilio (reported as Odiellus)palpinalis from Poland (Haitlinger 1987) . South -cott (1992) suggested that some of the specimen sidentified by Beron and Haitlinger were Leptusholmiae Southcott .

Mites reported as Leptus phalangii have bee nreported by Pfeifer (1956) and Evans (1910) onPhalangiidae (Phalangium opilio) in Berlin, Ger -many, and Midlothian, Scotland, respectively.Spoek (1964) also recorded this mite to be par-asitic on harvestmen from the Netherlands . Noneof these mites can be accurately identified at pres -ent and are best referred in Table 1 as Leptus sp .Meade (1855) reported "harvest-men" from En -gland were frequently infested by a bright redparasitic mite, which he identified as Trombi -

from Phalangiidae (M. morio) on Iceland . He notonly recorded both mites from a single specie sof harvestmen, but in two cases he recorded whathe felt were these species from individual har-vestmen . Although these cannot be identified wit hcertainty at this time, they are probably L. hol-miae. Until specimens can be studied I am re-ferring to them in Table 1 as Leptus sp.

Numerous mites from opilions in my collec-tion represent new records and include : Sclero-somatidae: Eumesosoma roeweri (Goodnight &Goodnight) from Alma, Nebraska, USA.; Krusasp . from 10 mi . W. Aquismon, San Luis Potosi ,Mexico ; Leiobunum aldrichi Weed from Tish-omingo State Park, Tishomingo County, Missis-sippi, USA ; Leiobunum flavum Banks fromBeaver's Bend State Park, McCurtain County,Oklahoma ; Merrymount Campground, 18 milesSW Nashville, Davidson, Tennessee, USA ;Leiobunum montanum montanum Suzuki fromMt. Ischizuchi, 1490-1745 m ., Ehime Prefecture ,Japan ; Leiobunum sp. from 2 km. N. Tasquillo ,Rio Tula, Hidalgo, Mexico; Leiobunum sp . nr .depressum Davis from 7 .5 miles S . George West ,Live Oak County, Texas, USA.; Leiobunumtownsendi from East Turkey Creek, ChiracahuaMountains, Cochise County, Arizona; outsid eHidden Cave (reared to deutonymph) and Her-mit Cave, Eddy County, New Mexico, USA . ;Leiobunum vittatum (Say) from Homesville ,Nebraska, USA . ; Trachyrhinus marmoratu sBanks from Pecos River, east of Pecos, PecosCounty, and Indio Mountains, 25 km S VanHorn, Hudspeth County, Texas, USA. Phalan-giidae : Odiellus pictus from Garland ; PenobscotCounty, Maine, USA . ; Phalangium opilio fromBowdoinham, near Cathance River, SagahahocCounty, Maine, USA.

Additional mites from my collection have bee nidentified by W . Calvin Welboum as Leptus spp .1-11 . They are as follows: Leptus sp. 1 is knownfrom a Sclerosomatidae (Leiobunum townsendi)and a Protolophidae (Protolophus singularisBanks) from Fort Bayard, Grant County, Ne wMexico, USA.Leptus sp . 2 is known from severalspecies of Sclerosomatidae : Eumesosoma roew-eri from 14 miles E. Burnet, Burnet County, Tex-as ; 7 .5 miles ESE Bandera, Bandera County, Tex-

dium phalangii (=Leptus phalangii) . He further as ; Texas Tech University Center, Junction,specified that the mite occurred on Leiobunumrotundum.

Sellnick (1940) recorded both Achorolophusignotus and Leptus (reported elsewhere in thepaper as Erythraeus) phalangioides (De Geer)

Kimble County, Texas, USA; Leiobunumflavu mBanks from Graham Creek, 5 miles SSE Zavalla ,Angelina County, Texas, USA ; Leiobunumtownsendi from Montague County, Texas, USA.Leptus sp . 3 is known from two species of Scle-

134

THE JOURNAL .OF ARACHNOLOGY

rosomatidae (Leiobunum aldrichi, L . nigripes gularis from England. Southcott (1992) statedWeed) from the W. Bank of J . Percy Priest Lake, that he felt some additional specimens reportedElm Hills Park, Davidson County, Tennessee, in the literature might be this species but that h eUSA. Leptus sp . 4 is known from a Phalangiidae could not be certain because he had not studied(Egaenus convexus) from Burgenland, Ruster any samples of the series reported. These ques-Hugelland, Austria . Leptus sp . 5 is known from tionable records are Phalangiidae : Mitopus mo-a Sclerosomatidae (Trachyrhinus marmoratus rio and Opilio ruzickai from Bulgaria (BeronBanks) from 39.6 miles SW Marfa, Presidio 1975); and Phalangium opilio, R . triangularis ,County, Texas, USA . Leptus sp . 6 is known from Lophopilio (reported as Odiellus) palpinalis froma Sclerosomatidae [Marthana nigerrima (Mull- Poland (Haitlinger 1987) .er)i from Tuba Mountains, S. Palawan Cabar, Leptus indianensis Fain et al . (1987) was de-Palawan, Philippines . Leptus sp . 7 is known from scribed from larvae collected on several specie sa Sclerosomatidae (Eumesosoma? sp.) from Joya of Sclerosomatidae : Leiobunum aldrichi (report-de Juan Mesa (outside), near La Laguna ; Ta- ed as L. longipes) and Leiobunum calcar (Wood)maulipas, Mexico . Leptus sp. 8 is known from a from 2 miles northwest Brazil, Clay County, In -Sclerosomatidae (Leiobunum sp .) from km 120 diana, USA . ; Leiobunum sp ., L. nigripes, L. spe-marker on Highway 70, San Luis Potosi, Mexico . ciosum Banks and L. ventricosum from 9 mile sLeptus sp . 9 is known from a Sclerosomatidae southwest of Crawfordsville, Montgomer y(Leiobunum sp .) from roadcut at Gomez Farias, County, Indiana, USA . New records from myTamaulipas, Mexico . Leptus sp . 10 is known from collection include L. nigripes Weed from 4 mile sa Sclerosomatidae (Lacinius ephippiatus) from ESE Morris on Pine Bluff Road, Grundy County,Wr. Wald, Latisberg (Cobenze) E-Mg ca . 380– Illinois, USA . ; L. formosum (Wood) from Po -400 m, Wien XIX, Austria. Leptus sp. 11 is known tomac River and Chesapeake Bay junction ,from a Phalangiidae (Mitopus morio) from Wr. Wakefield, Virginia, USA .Wald, Rekawiokel, near Bidf., N. G., Austria .

Leptus jocquei Fain & Eisen (1987) was de -Leptus bicristatus Fain & Eisen (1987) was described from nine larvae taken from "Opilions "

scribed from a larva on an "Opilion" from Cho- collected in Dembo, Plateau Nyika, Malawi (5 –wo Rocks, Plateau de Nyika, Malawi (6–8 De- 20 December 1981) . The host has now been iden-cember 1981) . The host has now been identified tified as a Phalangiidae, Cristina lettowi (Kauri ,as a Phalangiidae, Cristina lettowi (Roewer) pers . commun .).(Kauri, pers. commun.) .

Leptus kalaallus Southcott (1992) is thus farLeptus gagrellae (Oudemans 1910b) was orig- known only from the Phalangiidae, Mitopus mo-

inally described in combination with Achorolo- rio, collected in Greenland . The larval mites werephus Berlese . It was described from a Scleroso- found attached to the opilion abdomens .matidae (Gagrella sp .) from Tjibodas, West Java

Leptus lomani (Oudemans 1903b) was de-Prov ., Indonesia . This species was redescribed scribed from a Gonyleptidae, Lycomedicus (re-and illustrated by Oudemans (1913) .

ported Discocyrtus) funestus (Butler), from Chile .Leptus hidakai Kawashima (1958) was de- This species was redescribed and illustrated by

scribed from larvae collected on a Clubionidae Oudemans (1913) . The original series of 10 mite sspider (Chiracanthium sp .) and a Phalangiidae was reported to have been collected by J . C. C.(Opilio pentaspinulatus Suzuki) . All specimens Loman in 1900 . Other sources indicate that Janwere obtained on 24 June at Tachibana-yama, C . C. Loman, of Amsterdam, did not collect theKasuya-gun, Fukuoka Prefecture, Japan . A har- specimens himself. The harvestmen were prob-vestmen is illustrated in the original description ably collected by Prof. Dr. Ludwig Plate and for-with eight mites attached to its legs and abdo- warded to Oudemans by Loman . The only ex-men .

arnples of this harvestman reported in th eLeptus holmiae Southcott (1992) is a wide-rang- literature from Chile during the same time perio d

ing species in the Holarctic region . It is recorded was by Loman (Cekalovic K. 1985) . Loman(Southcott 1992) from a free living-example col- (1899) stated that there were several specimen slected in the Burzyanskij region, Bashkir ASSR ; ofL. funestus from Corral that were in the Plateand as ectoparasites on Phalangiidae : Mitopus collection . Therefore, I am herein restricting themorio from Denmark, Iceland, Poland, Sweden ; type locality of L. lomani to Corral (39°53'S ,Opilio sp . from Sweden ; Opilio canestrinii (Tho- 73°25'W), Valdivia, Chile .rell) from Denmark; Phalangium opilio from En-

Leptus nearcticus Fain et al. (1987) was de -gland; Rileana (reported as Platybunus) trian- scribed from larvae collected off three species of


13 5

Sclerosomatidae : Leiobunum aldrichi (reported

ACKNOWLEDGMENT Sas L. longipes), L . nigripes and L. vittatum from Like any project which is world-wide in scope ,2 miles northwest Brazil, Clay Co ., Indiana (1— many individuals were called upon for their as -18 September 1986). Fain et al . (1987) reported sistance. My colleagues who sent reprints of thei rother samples from the type locality from Leiob- publications are gratefully acknowledged. Dr . Janunum sp . (females) . These have now been iden- Buchar (University Karlovy, Praha), Dr . Asho ktified as L. aldrichi .

A. Hooli (Karnatak Science College, Dharwad) ,Leptus oudemansi (Karppinen 1958) was orig- Dr . Plamen G . Mitov (University of Sofia "Kli-

inal proposed as a replacement name in the genus ment Ohridski", Sofia), Mr . Sergio Sanchez-PenaAchorolophus . This name was provided because (Texas Tech University, Lubbock), and Mr . Lou -Achorolophus gracilipes Oudemans 1910a, was is M. Sorkin (American Museum of Natural His -preoccupied by Rhyncholophus gracilipes Kra- tory, New York) kindly supplied additional use-

mer 1897 ; both were considered by Karppinen ful literature . Dr . Howard E . Evans (Colorad o

(1958) to belong to Achorolophus . Both are now State University, Fort Collins) is thanked for hi s

considered by Southcott (1992) to belong to Lep- useful comments on pompilid wasps . Dr. Robert

tus and are thus still homonyms . Oudemans' G . Holmberg (Athabasca University, Athabasca )

(1910a) original specimens were found on a Cos- kindly provided me data on the numerous spec -

his(Cynorta sp .) from Surinam . This spe- imens ofLeptus spp. obtained from harvestmen

metidae his collection. Dr. Francis G . Howarth (Bisho pcies was redescribed and illustrated by Oude- Museum, Honolulu) is thanked for his corn -mans (1913) .

ments on harvestmen-like spiders of Hawaii . Dr .Leptus puylaerti Fain & Eisen (1987) is known Richard A

. Humber (USDA-ARS, Ithaca) i sby five larvae found attached to "Opilions" col -thanked for his useful comments on the manu -lected at Chowo Rocks, Nyika Plateau, Malawi

(6—18 Dec . 1981) . The host has now been iden- script (especially the fungi section) and for send-

tified as a Phalangiidae, Cristina lettowi (Kauri, ing copies of several difficult to obtain papers.

pers . commun .) .

Dr . Hans Kauri (Museum of Zoology, Universit y

Leptus polythrix Fain & Eisen (1987) is known of Bergen, Bergen) is thanked for his identifica-

by eight larvae found attached to "Opilions" col- tions of the opilion hosts of Leptus spp . fromlected at Dembo, Nyika Plateau, Malawi (5—20 Malawi . These harvestmen were loaned to Dr .December 1981) . The host has now been iden- Kauri by the Musee Royal de l'Afrique Centrale ,tified as a Phalangiidae, Cristina lettowi (Kauri, Tervuren. I thank Dr. Plamen G. Mitov for hispers . commun.) .

comments on parasites of harvestmen in Bul -Leptus stieglmayri (Oudemans 1905) was de- garia and for his permission to publish his man y

scribed from Opiliones collected in Santa Cruz, new records . Dr . Nobuo Tsurusaki (Tottori Uni -Rio Grande do SW, Brazil . Oudemans (1913) versity, Tottori) is thanked for his comments o nredescribed this species and recorded a specimen gregarine and fly parasites in Japan, his aid i nthat was collected from a beetle (Cleridae) col- providing difficult to obtain literature and fo rlected in Brazil .

translating some papers in Japanese . Mr . W. Cal -A probable new genus (near Leptus) is under vin Welbourn (Acarology Laboratory, The Ohi o

study by W . Calvin Welbourn . Thus far, mem- State University, Columbus) is thanked for hi sbers are only known from harvestmen from my many identifications of mites . Dr . George O .collection obtained in Chile . The new specimens Poinar, Jr. (University of California, Berkeley)are known from two species of Neopilionidae is thanked for his useful suggestions during th e(Thrasychirus modestus Simon, Thrasychirus preparation of this manuscript and for his com-dentichelis Simon) from Isla Deceit Caleta To- ments on the manuscript . I thank Dr . Normanleda, archipielago Cabo de Hornos, Magallanes, V . Homer (Midwestern State University) for hisChile . This is the southern most record for a many kindnesses shown me during the prepa-harvestman parasite . Other host records include ration of the manuscript . He helped me obtainspecies of Gonyleptidae : Eubalta meridionalis some literature and assisted with the adminis -(Sorensen) from Reserva Forestal Magallanes, trative details in obtaining funds for publication .8 km west Punta Arenas, Magallanes, Chile ;Metagyndes pulchella (Loman) ; Niebla, near More importantly, almost two decades ago Dr .Valdivia, Chile ; Acanthoprocta pustulata Loman Homer served as my professor of bacteriology,from Cerro Nielol, Temuco, Chile .

parasitology and arachnology .

136


Table 1 .-List of pathogens and parasites grouped by opilion host.

Host Parasite Source

Family? incertae sedisharvestmen, England ?Leptus sp. Evans 1910 ; Savory 1938harvestmen, Netherlands Leptus sp. Spoek 196 4Opliones, Brazil Leptus stieglmayri Oudemans 190 5Opiliones, Mexico Erythraeidae Martinez Crespo and

Morales Soto 1979Opiliones, U .S .A Leptus spp. Welbourn 198 3Opilionide, Israel Allothrombium Feider 197 7

chanaanenseopilionids Leptus sp . Mullen 198 8phalangid, U .S .A ?Leptus sp . Cox et al . 192 1Phalangium crassum, Sciadiophora Leger 1897

France phalangii

Suborder LaniatoresFamily Triaenonychidae and/o r

Synthetonychidaegen . sp., New Zealand Acarina, gen . sp. Forster 195 4

Chalcidae, gen . sp . Forster 195 4Nematoda, gen . sp. Forster 195 4

Family TriaenonychidaeEquitius doriae Acarina, gen . sp. Hunt 197 9

Gregarinasina, gen . sp. Hunt 1979

Family Assamida eOppalnia sp ., India

Family Gonyleptidaegen . sp . Brazil

Acanthoprocta pustulataDiscocyrtus invalidu sEubalta meridionalisGonyleptes fragilisLycomedicus funestusMetagyndes pulchellaZygopachylus albomarginis

Family CosmetidaeCynorta sp .Paecilaemana quadripunctaVonones sayi

Suborder CyphopalpatoresSuperfamily TroguloideaFamily Troguloida e

Trogulus tricarinatus

Family NemastomatidaeCarinostoma ornatu mParanemastoma radewi

Arachnocystisarachnoidea

Contospora opalniae

Echinoocysta phalangi i

Torrubiellagonylepticida

N . gn . nr. Leptus sp .Cyclorrhapha, gen. sp .N . gn. nr. Leptus sp.Agamomermis sp .Leptus loman iN . gn . nr. Leptus sp .Eumycota, gen . sp .

Leptus oudemans iAgamomermis sp.Acarina, gen . sp .Cosmetophilus vonones

Doliospora troguli

Gregarinasina, gen. sp .Gregarinasina, gen . sp.

Devdhar 1962 ; Devdha rand Gourishankar 197 1

Devdhar 1962 ; Devdharand Amoji 1978 a

Amoji and Devdhar 197 9

Moller 190 1

herei nSoares 194 5hereinMattheisen 1974Oudemans 1903bhereinMora 1987

Oudemans 1910aPoinar 198 5herei nCokendolpher 199 1

Geus 1969

hereinherei n


13 7

Table 1 .—Continued .


Agamomermis sp . hereinPyza bosnica Gregarinasina, gen . sp . herei n

Superfamily PhalangioideaFamily Neopilionida e

Thrasychirus dentichelis N . gn . nr. Leptus sp. herei nThrasychirus modestus N . gn . nr . Leptus sp. herein

Family ProtolophidaeProtolophus sp ., U .S .A . Agamomermis sp . Poinar 198 5Protolophus singularis Leptus sp . #1 herei n

Family SclerosomatidaeMetopilio grou p

Globipes sp . Acarina, gen . sp. herein

Subfamily LeiobuninaeEumesosoma? sp . Leptus sp. #7 hereinEumesosoma roeweri Leptus sp . herein

Leptus sp . #2 hereinLeiobunum sp ., U .S .A . Leptus indianensis Fain et al. 198 7Leiobunum sp . near Leptus sp. herei n

depressum, U.S .A .Leiobunum sp., Hidalgo, Leptus sp . herein

MexicoLeiobunum sp ., San Luis Leptus sp . #8 herei n

Potosi, MexicoLeiobunum sp ., Leptus sp . #9 herein

Tamaulipas, MexicoLeiobunum aldrichi Leptus sp . herei n

(=Leiobunum longipes) Leptus sp . #3 herei nLeptus indianensis Fain et al. 198 7Leptus nearcticus Fain et al . 198 7

Leiobunum blackwalli Anthorhynchus Pfeifer 1956(=L. hassiae) sophiae

Entomophaga batkoi Balazy 197 8Erythraeidae, gen . sp. Sankey 1949 b

Leiobunum calcar Leptus indianensis Fain et al . 198 7Leiobunum globosum Agamomermis sp . Tsurusaki 198 6

Gregarinasina, gen . sp . Tsurusaki 198 6Leiobunum flavum Leptus sp. herein

Leptus sp . #2 hereinLeiobunum formosum Leptus indianensis herei nLeiobunum manubriatum Agamomermis sp . Tsurusaki 198 6

Gregarinasina, gen. sp . Tsurusaki 1986Leiobunum montanum Leptus sp . herein

montanu mLeiobunum nigripes Leptus sp. #3 herei n

Leptus indianensis Fain et al . 1987 ; herei nLeptus nearcticus Fain et al . 198 7

Leiobunum politum Gregarinasina, gen . sp. hereinLeiobunum rotundum Anthorhynchus Ormieres and Baudoin 197 3

longisporaAnthorhynchus Pfeifer 1956

sophia eDoliospora repelini Ormii res and Baudoin 1973

138

THE JOURNAL OF ARACHNOLOG Y


Host Parasite Sourc e

Entomophaga batkoi Batazy 197 8Erythraeidae, gen. sp . Sankey 1949 bLeptus sp . Meade 185 5Sciadiophora Pfeifer 1956

phalangiiLeiobunum rumelicum Gregarinasina, gen . sp. hereinLeiobunum speciosum Leptus indianensis Fain et al. 1987Leiobunum townsendi Acarina, gen . sp. Elliott and Reddell 1987;

herei nLeptus sp. herei nLeptus sp. #1 hereinLeptus sp . #2 herein

Leiobunum ventricosum Acarina, gen . sp. hereinLeptus indianensis Fain et al. 198 7

Leiobunum vittatum Leptus sp. hereinLeptus nearcticus Fain et al . 198 7Nomuraea atypicola Greenstone et al . 198 8

Nelima sp ., Japan Trombidium sp. Yokogawa 194 0Nelima nigricoxa Forcipomyiinae, herei n

gen . sp .Nelima paessleri Acarina, gen . sp . hereinNelima silvatica Erythraeidae, gen . sp . Sankey 1949 bTogwoteeus biceps Agamomermis sp. Poinar 198 5

Leptus sp. herei n

Subfamily GagrellinaeGagrella sp ., Indonesia Leptus gagrellae Oudemans 1910bGagrellopsis nodulifera Acarina, gen . sp. herei nGagrellula ferruginea Forcipomyiinae, herei n

gen . sp.Gagrellula saddlana Sciadiophora Devdhar 1962 ; Devdhar

gagrellula and Amoji 1978bKrusa sp., Mexico Leptus sp . hereinMarthana nigerrima Leptus sp. #6 herei nMetagagrella tenuipes Charletonia southcotti Kawashima 196 1Prionostemma panama Acarina, gen . sp. hereinTrachyrhinus marmoratus Leptus sp . herein

Leptus sp. #5 herei nTrachyrhinus rectipalpus Acarina, gen. sp . herein

Family Phalangiidaegen . sp., England Hymenostilbe Leatherdale 1970

verrucosaPandora phalangicida Leatherdale 1970

gen. sp ., Germany Sciadiophora caudata Rossler 188 2Sciadiophora fissidens Rossler 1882

gen . sp., Sweden Pandora phalangicida Lagerheim 189 8

Subfamily PhalangiinaeBunochelis canariana Charletonia enghoffi Southcott 199 1Cristina lettowi Leptus bicristatus Fain & Elsen 198 7

Leptus jocquei Fain & Elsen 1987Leptus polythrix Fain & Elsen 1987Leptus puylaerti Fain and Elsen 198 7

Phalangium sp ., Italy Allothrombium Oudemans 1910 aneapolitum


139



Phalangium sp ., Sweden Leptus sp . De Geer 177 8Phalangium sp., Europe Sciadiophora Geus 1969

phalangiiPhalangium opilio Agamomermis sp . Pfeifer 1956; herein

(=Phalangium cornutum) Agamomermis Haldeman 185 1phalangii

Agamomermis Rudolphi 181 9truncatula

Anthorhynchus Schneider 1887; Ormieressophiae and Baudoin 1973

Brachylecithum sp . Gabrion and Ormieres 197 3Digenea, gen . sp . Creplin 1846Doliospora repelini Leger 1897; Ormiere s

and Baudoin 197 3Erythraeidae, gen . sp . Sankey 1949bGregarinasina, gen . sp. hereinHeterorhabditis Poinar and Thomas 198 5

heliothidisLeptus sp . Evans 1910 ; Pfeifer 1956;

Abro 1988, 1991 ; hereinLeptus holmiae Southcott 1992Leptus holmiae? Haitlinger I98 7Pandora phalangicida Ellis 1956; Leatherdale

1958, 197 0Pseudhymenolepis Gabrion 197 7

redonicaRhabditida, gen . sp. Pfeifer 195 6Sciadiophora caudata Rossler 1882; Ormiere s

and Baudoin 1973Sciadiophora fissidens Rossler 1882Sciadiophora Johansen 189 4

geronowitschiSciadiophora phalangii Leger 1897 ; Geus 1969Steinernema Poinar and Thomas 1985

carpocapsaeXenorhabdus Poinar and Thomas 198 5

luminescensXenorhabdus Poinar and Thomas 198 5

nematophilusRilaena triangularis Anthorhynchus Pfeifer 195 6

(=Platybunus triangularis) sophiaeErythraeidae, gen . sp. Sankey 1949bLeptus holmiae Southcott 199 2Leptus holmiae? Haitlinger 198 7Nematoda, gen. sp . Sankey 1949aSciadiophora phalangii Pfeifer 195 6

Zacheus anatolicus Gregarinasina, gen. sp . hereinZacheus crista Agamomermis sp. herein

Allothrombium hereinneapolitum

Gregarinasina, gen . sp . hereinZacheus hebraicus Marina, gen . sp. herein

Subfamily Oligolophina eLacinius ephippiatus Agamomermis sp. herein

140



Host

Parasite

Source

Pfeifer 195 6

hereinhereinhereinhereinSilhavy 196 1hereinhereinPfeifer 195 6Burton and Burton 1984Stipperger 1928; hereinOrmieres and Baudoin 197 3

Pfeifer 195 6

Ormieres and Baudoin 1973Sankey 1949bhereinStipperger 1928 ; Evans

et al . 1961 ; Abro 1988 ;herein

hereinSouthcott 1992Sellnick 1940 ; Berea 197 5Southcott 1992Rossler 1882 ; Ormieres

and Baudoin 197 3Stipperger 1928 ; Pfeifer

1956 ; Ormieres andBaudoin 197 3

Ormieres and Baudoin 197 3

hereinEvans 1948hereinhereinRossler 1882 ; Ormieres

and Baudoin 197 3Sankey 1949bOrmieres and Baudoin 197 3

Ormieres and Baudoin 197 3Balazy 1978 ; Keller 198 7Sankey 1949bhereinSankey 1949bherein

Lacinius dentiger

Lacinius horridus

Mitopus morio(=Opilio grossipes)

Mitopus sp., France

Odiellus lendliOdiellus pictus

Odiellus spinosus(=Odius spinosus)

Oligolophus hanseni iOligolophus tridens

Paroligolophus agrestis

Anthorhynchussophia e

Gregarinasina, gen . sp.Leptus sp. #1 0Agamomermis sp .Gregarinasina, gen . sp.Sciadiophora phalangi iAgamomermis sp .Gregarinasina, gen. sp .Rhabditida, gen . sp.Acarina, gen . sp.Agamomermis sp.Anthorhynchus

longisporaAnthorhynchus

sophia eDoliospora repeliniErythraeidae, gen . sp .Gregarinasina, gen . sp .Leptus sp .

Leptus sp. #1 1Leptus holmiaeLeptus holmiae?Leptus kalaallusSciadiophora caudat a

Sciadiophora phalangii

Sciadiophoraclaviformis

Gregarinasina, gen. sp .Anoplius marginatusGregarinasina, gen . sp.Leptus sp .Sciadiophora caudata

Erythraeidae, gen . sp .Anthorhynchus

sophiaeDoliospora repeliniEntomophaga batkoiErythraeidae, gen . sp.Leptus sp .Erythraeidae, gen. sp .Leptus sp.

Subfamily Opilionina eEgaenus convexus Leptus sp . #4 herein

Trombidium hungarium Kobulej 195 7Opilio sp., Europe Agamomermis sp . Kastner 192 8

Agamomermis Rudolphi 181 9truncatula


14 1

Table 1 . —Continued .


Leptus holmiae Southcott 199 2Opilio canestrinii Leptus holmiae Southcott 199 2Opilio dinaricus Gregarinasina, gen. sp . hereinOpilio parietinus Agamomermis sp. Unzicker and Rotramel 197 0

Anthorhynchus Ormieres and Baudoin 197 3longispora

Doliospora repelini Ormieres and Baudoin 197 3Erythraeidae, gen . sp. Sankey 1949bLeptus sp . Evans et al. 196 1Microsporidium Silhavy 1960

weiseriSciadiophora phalangii Pfeifer 1956

Opilio pentaspinulatus Leptus hidakai Kawashima 195 8Opilio ruzickai Gregarinasina, gen . sp. herei n

Leptus holmiae? Beron 197 5Opilio saxatilis Gregarinasina, gen. sp . herein

Subfamily PlatybuninaeLophopilio palpinalis Agamomermis sp . herei n

(=Odiellus palpinalis)Gregarinasina, gen . sp . hereinLeptus sp . Haitlinger 198 7Leptus holmiae? Haitlinger 198 7Sciadiophora frssidens Rossler 1882

Megabunus diadema Actinocephalus Ormieres and Baudoin 197 3megabuni

Doliospora repelini Ormieres and Baudoin 197 3Platybunus bucephalus Anthorhynchus Ormieres and Baudoin 197 3

longisporaDoliospora repelini Ormieres and Baudoin 197 3Sciadiophora phalangii Ormieres and Baudoin 197 3

Platybunus pinetorum Sciadiophora phalangii Pfeifer 1956

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Manuscript received 4 August 1992, revised 10 Marc h1993 .

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