162 Plant Protec ti on Quarterl y VoJ.6(4) 1991 not been seen, and that Tucker had sug­ Taxonomic background of the redlegged earth gested that H. des tructor was parthenoge­ netic. The situation was not helped when Halotydeus destructor (Tucker) (Acarina: ) the illustration of the male in Newman (1 925b and elsewhere) was reprod'uced in R . B. H alliday, CSIRO Division of Entom o logy, G PO Box 1700, Canbe rra, Newman (l936) labell ed as a female. ACT 2601, Australia . Meyer and Ry ke (1 960) and Meyer (1 981) also sta ted that the males were not known, and attributed to Tucker the view Summary then realised that the RLEM of Western that the species is probably parthenoge­ The early literature on red legged ea rth Australi a was not the sa me as Froggatt's netic. mite Hniotydell s des tnlctor (Tucker) species from New South Wales, and be­ Tucker's (1 925) observation of "several contains man y bib liographic and gan referring to the former as Pen thalells cases of apparen t parthenogenesis" could n om enclatural errors, w hich h inder deslrllctor (Jack) (Newman 1925a, 1925b). refer to complete female-female parthe­ study of its taxonom y and biology. Tucker (1925) described the earth fl ea nogenesis, or to the production of males Th ose errors are here corrected. It ap­ taxonomica ll y as P. destructor, and gave it from unfertilized eggs, a well-known pears as if the species occurs only in the new common name of black sand phenomenon in many mite groups. southern Africa, Aus tralia, and New mite. He also stated that Jack (1908) had Tucker also sta ted that copulati on had Zealand. Research is required to deter­ not given a description of the earth fl ea, never been observed. This in itself is not m ine whether it also occurs in Mediter­ but had referred to it as P. des tructor. surpri sing, since many practise ranean Europe, w h ere several rela ted Tucker is mistaken on both counts - Jack non-copulatory sperm transfer, in which species occur. Research is also required did describe the general appea rance of the the males deposit a spermatophore which to allow the confident taxonom ic recog­ spec ies, but d id not use the name destruc­ is later picked up by a female. Solomon nition of H. destmctor, so that it may be tor, or any other scienti fic na me. Tucker's (1937) described the web-spinning behav­ distinguished from related species in statements appear to have been based on iour of H. dest ructor, and noted that there Australia and elsewhere. Possible fu ture some manuscript of Jack's that was never were "small globules of water-like fluid biological control strategies cannot be published . His error of attributing the spaced at intervals along the threads". He fully exploited until these questi ons are name destructor to Jack was repea ted by observed that the specimens spinning answered. subsequent authors (e.g., Womersley web were smaller than adult fem ales, and 1933, 1941 , Newman 1925a. 1925b, Thor were probabl y males. It was suggested at Introduction and Will mann 1941 , Strandtmann and the time that the webbing contains, or is The redlegged earth mite Halolydell s de ­ Tilbrook 1968). Newman (1925a, 1925b) composed of, spermatophores, perhaps stnletor was described taxonomically by compounded the confusion by referring with sperma tozoa in the fluid droplets Tucker (1925), under the name Pel/lilalells to a publica tion by "W. E. Jacker", which (K. R. Norris, in correspondence, 1938). destructor. This bland sta tement conceals keyed and described P. des/rllc/or. This These sperma tophores are remarkably the fa ct that the species has had an unusu­ mythical author appears to be a chimera different fro m those typically produced all y interesting and compli ca ted of R. W. Jac k and R. W. E. Tucker. The by prostigmatid mites. These are usuall y nomenclatural and bibliographic history. publication referred to is probably Tucker erect mushroom-like structures of distinc­ The early literature on the species is full of (1925), since Newman quotes it as using ti ve shape, produced Singly, in which a errors, misunderstandings, and misquo­ the name black sa nd mite, which had packet of sperm is supported on a sta lk, tations. These erro rs confound attempts to been introduced by Tucker. Jack eventu­ rather like a golf ball on a tee. The sta lk determine the native range of the spec ies all y settled the issue by him self attribut­ often has attached ramifica ti ons and ex­ and its present geographic range, and to ing the name destrllctor to Tucker Oack, crescences that are of consistent shape consolidate the avaiJable information on 1942). within a species (see for example Krantz its biology. It is therefore timely to sum­ Swan (1 934) perfo rm ed the very useful 1978, Figure 12). Preliminary observa tions mari ze some of the history of the species, service of correcting a number of have shown that the specimens of RLEM in an attempt to clarify these questions. nomencl atura l and hi storical errors that that spin web never contain eggs, and had been made by previous authors. have an interna l organ resembling an The early literature on redlegged These incl uded some spelling erro rs, aedeagus, w hile specimens containing earth m ite which should be disregarded eggs do not spin web and do not have this The redlegged earth mite (RL EM) first Hn /o tydnells, Haiodytaeus, Notophal ll s, bi­ structure 0. C. Otto, personal communi­ appeared in the litera ture in 1908 under cololl r. dicolor. Newman (1936) then acted cation 1991). Also, the webbing produced the name earth fl ea (Ja ck 1908). Jac k gave to co rrec t his earlier erro rs, but Swan's ef­ by RLEM bears a striking resemblance to brief descriptions of its life history and bi­ forts appea r to have been overl ooked by that of Linopodes sp., in the related family ology, and an account of the damage tha t some other authors. , in which webbing been the species was causing to vegetables in shown to act as a carrier of sperma to­ the Cape Province of South Africa. He did T he existen ce of m ales phores (Ehrnsberger 1989). not, however, use any scientific name to There has been some doubt as to the mode Strandtmann and Ti lbrook (1 968) dis­ describe the species. Banks (1 915) referred of reproduction of RLEM , and w hether or tinguished the males and females of to the South African earth fl ea as an uni­ not males ex ist. Jac k {I 908) described the Hnlolydeus siglliensis by the morphology dentified species of Pen t1mlell s. Newman male as being small er than the fema le, of the genitalia and the number of genital (1 920) recognized the presence of earth and as having an abdomen that was ta­ seta e. Saker (1990) also described internal fl ea in Western Australia, and later (1923/ pering rather than rounded . Newman structures, notably the sperm sacs, that 1924) identified it as Nolophallils bieolor (l 925a, 1925b) published photographs la­ allow the recognition of males in other Froggatt 1921 , which had just been de­ belled "adult male", without ex planatory species of . There appears to sc ribed from New South Wales. At the tex t. Neither of these authors offered any be no doubt that males exist in RLEM , but same time Newman (1923) coi ned the evidence that their specimens were in fa ct the descriptions that have been published name red legged velvet ea rth mite for the males. Womersley (1933) confused the is­ to date are not adequate to allow them to Western Australi an species. Newman sue when he said that males of RLEM had be recognized morphologically. Plant Protection Quarterly VoI.6(4) 1991 163 Geographic distribution of and described it as bl ac k with red legs and are neither egregius nor destructor, but be­ H. desmlctor a medio-dorsal red spot. This red spot re­ long to another species of unknown iden­ Halotydeus destructor was described from sembles that of (Duges tity. The status of this record should be Cape Province, South Africa, where it is a 1834), but Berlese explicitly states that H. critically re-assessed on the basis of widespread and abundant pest. Its biol­ hydrodrortills has its anus terminal, while freshly coll ected specimens. Schuster ogy and behaviour in South Africa have the anus is dorsally placed in Penthalells . (1958) rightly pointed out that the red pig­ been reviewed by Meyer (1981). The first H. hydrodromus was subsequently found ments in these species are lost when the record of its occurrence in Australia was to be abundant on beach roc ks at low tide mites are preserved in alcohol or mounted at BunburyWA in 1917 (Newman 1925b). in Ireland (Halbert 1920). H. hydrodromlls on slides, so old museum specimens may Hearsay reports of the time suggested that variety albolineatus Halbert 1915 was de­ be difficult to identify. it came from ship's ballast of South Afri­ sc ribed from rocks on a beach in ueland, It should also be noted that H. destrllctor can origin (Johnson 1930). Once in Aus­ and is distinguished by a white dorsal varies in colour. Womersley (1933) noted tralia it appears to have spread very stripe. H. hydrodro11/11 S aiboiilleatus also oc­ the existence of specimens with a brown quickly, and was recorded in both South curs in the littoral zone on the western dorsal surface sharply demarcated from Australia and New South Wales in 1930 Mediterranean coast of France (Schuster the black ventral surface. It has subse­ (Johnson 1930, Anonymous 1930a). 1958). Hal otydells mol/is Lu xton 1986 was quently been shown that these specimens Newman (l925b) observed that H. de­ collected on a mangrove beach in Hong are those infected with a parasitic fungus structor had been found only in South Af­ Kong, H. signiellsis Strandtmann and (Petch 1940). Tucker (1925) noted some rica and Western Australia, and inferred Tilbrook 1968 is found in moss on Signy specim ens in South Africa that he de­ that South Africa was its country of ori ­ Island (So uth Orkney Islands), and Talker scribed as "greyish or yellow mottled" in gin. This view became established in later et al . (1981) recorded an unidentified spe­ colour, which may represent cases of the literature on the species, from Anony­ cies of Halotydells from the intertidal zone same fungal infection. mous (1930b) to Wall ace and Mahon in the Philippines. (1 971). It was later reported as being in­ The situation surrounding Halotydeus Family-level relationships troduced into New Zealand (Dumbleton, egregill s is confused. Berlese (1891) de­ Some authors have stated that H. destruc­ 1947), and as being present in Zimbabwe sc ribed Pentilaiells egregius from several tor is related to a group of predators (e.g., and Malawi (Jack 1942, Goldsmid 1962). localities in Italy (Venice, Tuscany, Na­ Tucke r 1925). This impression was cre­ Published maps summari zing these re­ ples, Sicily). Berlese then (1903) trans­ a ted by the use of a classification that sults show its distribution to be southern ferred P. egregi ll s to a new genus pla ced Ha lotydeus in a broadly conceived Africa, Australia, and New Zealand Chromotyda ell s Berlese 1903, on the family Eupodidae, which included (Anonymous 1958, Anonymo us 1960). grounds that its anus was ventral, and not groups of predatory mites that are now Macfadyen (1954) recorded one specimen dorsal as it is in Pellthnlel/ s. This is puz­ placed elsewhere. However, most mod­ of H. destrllctor from Ian Mayen Island, zling, since Berlese (1891) clearly illus­ em taxonomists consider that Halotydws Greenland (71°N), and suggested that it trates P. egregi ll s with a terminal anus, belongs to the separate family ma y have been accidentally introduced which would place it in Hnlotydell s (see for Penthaleidae within the superfamily there in food supplies. This remarkable example the key of Strandtmann 1981). Eupodoidea (for example Krantz 1978, record cannot be confirmed, and it is most Womersley (1941 ) suggested that P. Strandtmann 1981, Kethley 1982). In this unlikely that the species is actually estab­ egregills was actually a speci es of classification the genera have been di­ lished there (Wa ll ace and Mahon 1971 ). Haiotydells, and recorded it from Western vided among five smaller families, which Womersley (1933) raised the possibility Australia . Baker (1946) also considered more closely reflect the biology of their that RLEM had been introduced to South that H. egregius belonged in the genus constituent species. Africa from the Mediterranean climate Halotydeus, and recorded it from orchard The family are all preda­ areas of southern Europe. M.M.H . soil in Texas. Berlese (1891) and Thor and tory, and have strong chelicerae with op­ Wallace made several collecting trips to Willmann (1941) report that H. egregills is posable digits for biting prey. They occur Europe and north Africa between 1964 usually found on bare dry soil in the sum­ in soil, leaf litter, moss, under bark, and in and 1969 with the specifi c objective, mer, and is uncommon in wet habitats. caves. The family Strandtmanniidae co m­ among others, of determining whether H. The Australian records given by prises only two species, both of which oc­ dest ru ctor occurred th ere. He coll ec ted Womersley (1 941 ) do not appea r to be cur in leaf litter. Their feeding habits are great numbers of mites and Coll embola consistent with this observation, all hav­ unknown, but they have robust raptorial from pastures and other habitats in south­ ing been collected between April and No­ chelicerae that strongly suggest preda­ ern France, Spain, Morocco, and ltaly, but vember, or "in winter". Also, fi eld notes tion. The Eupodidae, Penthalodidae, and did not find H. destrll c/or (Wallace 1966). o f the time (by K.R. No rris) show that Penthaleidae have modified chelicera e However, his travels did not include the most of these specim ens were collected in with weak opposable digits, and al most eastern Mediterranean, on either the Afri­ moss, and not in pasture. They were re­ all appear to be plant feeders. The ca n or European shores. Jeppson et al. ferred to in these notes not as H. egregills, Eupodidae are found in damp soil, hu­ (1975) and Meyer(1 981) record H. destrllc­ but by an unpublished name that suggests mus, and moss, and have a worldwide tor from Cyprus, but this record requires that their legs are longer than those of H. distribution. Some species are co nfirmation, since it does not appear in lies/ru etor. It is unlikely that Womersley's fungivo ro us in commercia l mushroom the listings of the mites of Cyprus given record of H. egregius from Australia actu­ houses, others have been recorded feed­ by Georghiou (1959, 1977). 15ee 1I0te added all y refers to H. des tructor, since the colour ing on algae, but most have not been stud­ ill proof: page 1651 difference between these two species ied biologica lly. ThePenthalodidaeareall seems to be distinct, at least in living highly ornamented, and occur in moss, Other species of Halotydells specimens. The body of H. destructor has algae, and leaf litter. The species whose The genus Halotydws was erected by always been described as entirely black, biology has been studied are thought to Berlese (1891) to accommodate one spe­ while the red dorsal spot of H. egregills feed on algae, fungi, and li chens. cies, H. hydrodroltllls (Berlese and was described by Berlese, and has been The family Penthaleidae is a small one, Trouessa rt 1889), which was described observed in Australi an specim ens comprising only Halotydeus Berlese 1891, from rocks on the beach in France and (Womersley 1941). It is possible tha t Pellthaleus Koch 1836, Linopellthniells Italy. Berlese (1891) illustrated the species Womersley's specimens of "H. egregills" Willmann 1951, and Linopen thaJoides l64 Plant Protection Quarterly VoI.6(4) 1991 Strandtmann 1981. Womersley (1941 ) and ment. It will be necessa ry to distinguish wealth Institute of Entomology). Baker (1946) included the genera H. des tructor from o ther species of Baker, A.S. (1990). A survey of external Stereotydeus and Pelltlza[odes in the family Halotydws, notably H. egregills, H. morphology of mites of the superfamily Penthaleidae, but these are now consid­ hydrodromus, and H. hydrodromu s variety Eupodoidea Banks, 1894 (: ered to be better placed in the separate albolinentus. Conventional morphological ). Journal of Natural History family Penthalodidae. Linopell thnl ells characters should be sought, to allow 24, 1227-61. comprises one species collected in grass these species to be separated . During col­ Baker, E.W. (1946). New species of North and under rocks in the Austrian alps, and lecting. it is important that the collector and Central American mites of the fam · LillopentJUlioides comprises one species make detailed descriptions, illustrations, ily Penthaleidae (Acarina). JOllmal of the from a cave in New Zealand. Only two and preferably colour photographs, of Washingto" ACildemy of Sciences 36, 421 - species o f Penthaleidae ha ve been studied living and freshly-killed specimens, to 5. biologica lly, H. des tructor and P. major, record their colours. The co llector should Banks, N. (1915). The Acarina or mites. A and even these are not clearly defined also develop methods of long-term pres­ review of the group for the use of eco­ taxonomically. Many other names have ervation of specimens that allow their vi­ nomic entomologists. United States De­ been used in the genus Penthaleus, as re­ tal colours to be retained. It would also be partment ofAgric illture Report 108, 1-153. viewed by Thor and Willmann (1941), but useful to obtain fresh comparative mate­ Berlese, A. (1891). Acari, Myriopoda et most of these are no t taxonomically rec­ rial o f the species of Hal ol ydells that are Scorpiones hucusque in Italia repertn 60. ognizable, and their biology is completely known to occur in southeast Asia and Berlese, A. (1903). Acari nuovi. unknown. It is no t known whether any of North America . Manipulus [us. Redia 1, 235-52. these are economica lly important pests, Research is also required to determine Berlese, A. and Trouessart, E. (1889). Di­ but it is possible that some literature how male RLEM can be recognized mor­ agnoses d'acariens nouveaux ou peu records of P. major actually belong to dif­ phologically, and whether fertili 7.a tion is conn us. Bulletin de In Bibliotheqll e ferent species. necessary for completion of the life cycle. Scientifiqlle de l'Ollest 2, 121-43. If fertili za ti on is required for develop­ Dug~s, A. (1834). Recherches sur I'ordre Conclusions ment, it will be necessary for males and des Acariens. Annales des Sciences Despite its economic impo rtance. and de­ females to recognize each other, or for fe­ Natl

Note added in proof:

The Cyprus record of H. destructor is now known to be an error (L.R. Je ppson, personal communication). RLEM has not been recorded from Europe. R. B. Halliday