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The ground skink, Scincella lateralis, an unusual host for phoretic deutonymphs of a uropodine mite, Fuscuropoda marginata, with a review of analogous mite-host interactions.

Abstract.--Phoretic deutonymphs of the uropodine mite Fuscuropoda marginata (Koch) that usually attach to dung beetle hosts were found for the first time on a native ground skink, Scincella lateralis (Say), collected on the grounds of the Dallas Zoo, Dallas County, Texas. It is suggested that this mite and other acarine taxa may be preadapted to shift between arthropod hosts covered with chitinous sclerites and vertebrate hosts covered with keratinous scales.


Lizards are often and expectedly hosts to a variety of arthropods, including external mites (Leeuwenhoekiidae, Macronyssidae, Pterygosomatidae, Trombiculidae) or ticks (Argasidae, Ixodidae) and internal pentastomids (Cephalobaenidae, Porocephalidae, Sambonidae) (Wharton & Fuller 1952; Zumpt 1961; Yunker & Radovsky 1966; Hoogstraal & Aeschlimann 1982; Self 1982; Cruz 1984). A few less-well-known mites are associated with lizards in limited circumstances, internally (Trombiculidae, Macronyssidae) and externally (Erythraeidae, Paramegistidae) (Baker & Wharton 1952; Yunker & Radovsky 1966; Vercammen-Grandjean & Watkins 1965; Voss 1967).


On 27 April 1999, a native adult ground skink, Scincella lateralis (Say), was collected on the grounds of the Dallas Zoo, Dallas County, Texas. The collection site (32[degrees] 44.499' N, 96[degrees] 49.083' W) is a semi-wooded, unmanaged plot adjacent to the Bird and Reptile Building (Hartdegen et al. 2001). The skink was housed alone in a sealed, new, solid plastic container on new, dampened paper toweling until its examination two days later. At that time, the animal was found to be infested with numerous small, sessile mites that were not observed when it was collected. In all, 60 mites were collected with cotton-tipped swabs and preserved in 70% ethanol. Voucher specimens (Accession Number 229877) are deposited in the Parasitology Reference Collection at the National Veterinary Services Laboratories, Ames, Iowa.


Most of the mites were affixed to the animal's head (Fig. 1), although smaller groups were on the right forelimb and the basolateral tail. Because the skink was housed in a pristine container, the individual mites must have been present, though unattached, on the animal when it was collected; they became noticeable only after aggregation and affixation. On 28 July 1999, the mites were identified as phoretic deutonymphs of a species in the family Uropodidae and recognized as unusual on this host. In September, 1999, the mites were identified further as Fuscuropoda marginata (Koch) (c. f., Lehtinen, pers. comm.)


Uropodids, often called tortoise mites because of their general shape and leg articulations, compose one of several families of mesostigmate mites classified in the large and diverse acarine cohort Uropodina. The systematics and taxonomy of uropodines seem to be in a continual state of flux, with many opinions and little consensus among specialists on a stable arrangement and nomenclature (Johnston 1961; Evans & Till 1979; Hirschmann 1979; Farrier & Hennessey 1993). (Indeed, some workers would use for F. marginata the genus name Uroobovella, and some would use the family name Urodinychidae. In one reference (Dusbdbek & Bukva 1991), it is referred to under three different scientific names.)

Most uropodine mites (including those in the Uropodidae) are probably scavengers, detritivores, bacteriophages or fungivores, feeding on decaying organic matter or the microorganisms growing therein (Hughes 1959; Johnston 1982). However, the bionomics of relatively few species have been studied because few have any economic importance. One species, Uroactinia agitans (Banks), is a food competitor pest in the growth beds of earthworm producers (Stone & Ogles 1953), and a few species are known to prey on soft-bodied stages of small arthropods, free-living nematodes, or other invertebrates (Reid 1957; Johnston 1982).

Phoresy is a common behavior for uropodine mites, particularly so among the uropodids. One of the immature uropodid stages, the deutonymph, may occur in two types that differ in behavior and morphology (Faasch & Schaller 1966; Faasch 1967). One type is an undistinguished, sedentary developmental form (the so-called "dauernymph"), and the other is a distinctive form (the so-called "wandernymph") that is specialized for dispersal, primarily via more mobile or even volant arthropod hosts. Each phoretic deutonymph attaches to its host by means of an anally secreted fluid that solidifies in the air into a supportive, stem-like structure called a pedicel.

Fuscuropoda marginata is a relatively common and widespread species known from degraded stored food products and a variety of primarily edaphic habitats, especially in the Northern Hemisphere. It is the type species of the genus Fuscuropoda Vitzthum 1924, and it has been known probably at least since 1768, when what was possibly the same mite was described under the name Acarus vegetans De Geer. Hirschmann & Zirngiebl-Nicol (1962) and Hirschmann (1979; 1989) discussed some aspects of F. marginata systematics. Most published references to F. marginata are minor and incidental notes on its presence in surveys of soil arthropods (Ryke 1958; Krantz & Ainscough 1990; Farrier & Hennessey 1993). It occurs in both forest and grassland soils, in greenhouses, under stones and wood, in compost and decaying plant materials, numerously in moldy grain, and very commonly in chicken litter or other kinds of manure (Hughes 1948; Krantz 1961; Evans et al. 1961; Karg 1968; Jalil & Rodriguez 1970; Oswald & Minty 1971; Powell & S kaley 1975; Wisniewski & Hirschmann 1993). Only a few detailed works discuss its biology and morphology (O'Donnell & Axtell 1965; Faasch & Schaller 1966; Faasch 1967; Karg 1968; Jalil & Rodriguez 1970; Afifi 1983; Raut & Panigrahi 1991).

Although once thought to be primarily coprophagous or fungivorous (Evans et al. 1961), later studies have shown that F. marginata may feed sometimes on living plants (Karg 1968). However, it is largely predaceous on weak or immature mites and nematodes (Faasch 1967; Royce & Krantz 1991), on insect eggs and larvae (Jalil & Rodriguez 1970), or even on mollusks (Raut & Panigrahi 1991). Through its predatory activities, F. marginata may have some economic benefit in suppressing populations of filth flies around poultry facilities (O'Donnell & Axtell 1965; O'Donnell & Nelson 1967; Axtell & Rutz 1986) or in reducing plant damage by pestiferous slugs (Raut & Panigrahi 1991). On the other hand, Dhanakkodi (1990) found that F. marginata serves as an intermediate host for a sheep tapeworm, Moniezia benedeni (Moniez), in India, and Karg (1968) observed significant feeding damage caused by these mites on greenhouse cucumbers in Germany.

Only two previously documented instances were found of phoretic uropodids on lizards, both in the Australian Region and both on skinks. The first case (Domrow 1981) involved an undescribed species of Uroactinia infesting a delicate skink, Lampropholis delicata (De Vis), at Mount Molloy, Queensland, Australia, in April, 1978. The skink was found moribund and bearing "many hundreds" of phoretic deutonymphs. This mite was later redescribed and named U. domrowi Hirschmann (1990), but nothing more is known of it. The second case (Rowlands 1983) briefly cited and illustrated "large numbers" of unnamed uropodid deutonymphs attached around the limbs of a copper skink, Cyclodina aenea Girard, collected in November, 1981, from a garden area in the Milford area, Auckland, New Zealand.

Phoretic deutonymphs of F. marginata normally attach to various arthropods, particularly those that usually inhabit decaying organic materials and manure. Some recorded transport hosts include: beetles of the families Buprestidae, Carabidae, Coccinellidae, Hydrophilidae and Scarabaeidae (Muller 1860; Faasch & Schaller 1966; Faasch 1967), bibionid flies (Pecina 1970), millipedes (Faasch 1967) and sowbugs (Vitzthum 1943). In laboratory experimental choice studies (Faasch & Schaller 1966; Faasch 1967), phoretic F. marginata deutonymphs attached most readily to Geotrupes dung beetles, but in the absence of such hosts, they also attached to carabid beetles and millipedes. Indeed, F. marginata was less fastidious about attachment surfaces than was another common, similarly tested uropodid species, Uropoda orbicularis (Muller) (Faasch & Schaller 1966). Unlike the latter species, F. marginata wandernymphs would readily attach to smooth, water-repellent surfaces like the glass or plastic walls of the containers confin ing them. By contrast, U. orbicularis deutonymphs attached to the walls of the containers only when the surfaces were treated with the odor of their normal dung beetle transport hosts, suggesting that different species of uropodid wandernymphs may differ in the number and kind of cues necessary to elicit attachment behavior.

Such facultative acceptance of facsimile substrates for attachment probably accounts for the occasional, evidently accidental choice of lizards as transport hosts by some uropodid wandernymphs. The literature on F. marginata documents that manure-dwelling beetles are its usual hosts, but there are scattered, one-time records (Muller 1860; Vitzhum 1943; Pecina 1970) for a few other, uncharacteristic arthropod hosts. In addition to the two previously cited records for other species of uropodid deutonymphs attached to vertebrates (i.e., skinks), there is also a record of a Fuscuropoda sp. (probably F. marginata, in light of its abundance in chicken litter) attaching to broiler chicks in a poultry house (Rives & Barnes 1988). One might postulate that, occasionally, a local population of uropodids may produce a cadre of phoretic deutonymphs at a time or place where no or insufficient preferred hosts are present. Faasch (1967) found that declining quantity or quality of local suitable food is the trigger for produc tion of wandernymphs and their phoretic behavior. One can envision a number of scenarios in which wandernymphs might arise without access to suitable beetle transport hosts. For example, a relatively small quantity of manure (or other substrate) attracts a beetle bearing deutonymphs, some of which detach and colonize the site, even though the beetle finds that the site is insufficient for its needs and leaves without reproducing. By the time that the mites have completed a life cycle, the food resource is depleted, and new wandernymphs are produced. However, none of the usual transport host beetles are present, and the nymphs are left to die or to attach to whatever marginally acceptable animal may happen to move through the site. In the case of F. marginata, acceptability seems to be based primarily on the body surface of the transport host being covered by a smooth, relatively hard, hydrophobic substance, e.g., chitinous plates on arthropods, keratinous scales on lizards or even on the legs of birds. One ca n envision other scenarios, as well, but evidently, such circumstances do not occur frequently in nature, at least for those (possibly also infrequent) uropodid species with relatively few requirements for transport host acceptability.

In addition to the rarely documented instances where some phoretic uropodid species deutonymphs behaviorally stray from their normal arthropod hosts to lizard hosts, there are some recorded cases among the Acari where analogous and seemingly anomalous host shifts have occurred at the evolutionary/phylogenetic level. The Schizogyniidae is a small family of mestostigmatid mites whose members are usually associated with heavily sclerotized beetles, e.g., Passalidae, Carabidae, Scolytidae (Kinn 1966; Hunter & Rosario 1987). However, the unique species Indogynium lindbergi (Sellnick) is found only on burrowing snakes in India (Sellnick 1954). Another related mesostigmatid family, Paramegistidae, includes commensal mites associated with sclerotized arthropods, e.g., beetles, ants, myriopods (Tragardh 1906, 1907; Nickel & Elzinga 1970), plus species of the genus Ophiomegistus that occur on snakes and lizards in the East Indian Region (Voss 1967). Yet a third related mestostigmatid family, Diplogyniidae, includes mit es exclusively associated with beetles (Passalidae, Histeridae, Scarabaeidae), except for the unique species Ophiocelaeno selinicki Johnston & Fain found on snakes in the Solomon Islands (Johnston & Fain 1964).

Some instances of possible host shifts in the other direction, from vertebrates to arthropods, also are documented. The prostigmatid family Pterygosomatidae comprises nearly 100 species of parasitic mites that live on or under the scales of lizards (Hirst 1926; Davidson 1958; Bertrand et al. 2000), plus the genus Pimeliaphilus, whose few members are parasites of cockroaches, reduviid bugs or scorpions (Beer 1960; Newell & Rychman 1966). The hematophagous larval stages of the prostigmatid mite families Trombiculidae and Leeuwenhoekiidae (chiggers) normally feed on a variety of vertebrates, including many that feed on reptiles. However, each family contains one or two species whose larvae parasitize scorpions or millipedes (Andre 1943; Wharton & Fuller 1952; Audy 1956; Nadchatram & Dohany 1974).


Some or all of the preceding, putatively anomalous host-mite associations may have come about because the mites involved possess genetically preadapted host-selection behaviors triggered by a limited suite of acceptance cues, primary among which is a host integument consisting of smooth, hard, water-repellent scutes. The validity of this hypothesis would seem to be amenable to experimental verification.


We thank John H. Greve (Iowa State University, Ames) for putting the two authors in contact with one another and for supplying pertinent literature, and Pekka T. Lehtinen (University of Turku, Finland) for specifically identifying the mites and for supplying background information.


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