ORIBATID MTTES AND THE DECOMPOSITION OF PLANT TISSUES IN PALEOZOIC COAL-SWAMP FORESTS CONRAD C. LABA^a^EIRA, TOM L. PHILLIPS, AND RDY A. NORTON Made in United States of America Reprinted fmrn PALAIOS VoL 12. No. 4, A-agast, 1997 Copyright O 1997, SEPM (Socie^ for Sedimentary Geology) RESEARCH REPORTS 319 Oribatid M?tes and the Decomposition of Plant Tissues in Paleozoic Coal-Swamp Forests CONRAD C. LABANDEffiA Department ofPaleobhlogy, National Museum of Natural History, Smithsonian Institution, Washington, D.C. and Department of Entomology, University of Maryland, College Park, MD 20742 TOM L.PHILLIPS ^Department of Plant Biology, University oflUinois at Urbana-Champaign, Urbana, IL 61801 ROY A. NORTON ^Department of Environmental and Forest Biology, College of Environmental Science and Forestry, State University of New York, Syracuse, NY 13210 PALAIOS, 1997, V. 12. p. 319-353 Although oribatid mites are essential to the decomposition of plant tissues in modern temperate forests by assisting conversion of primary productivity to soil organic matter, little is known of their paleoecohgic history. Previously there has been scattered and anecdotal evidence document- ing oribatid mite detritivory in Pennsylvanian plant tis- sues. This study evaluates the incidence of oribatid mite damage for seven major coal-ball deposits from the Illinois and Appalachian sedimentary basins, representing a 17 million year interval from the Euramerican tropics. Al- though this interval contains the best anatomically pre- served plant tissues with oribatid mite borings in the fossil record, coeval oribatid mite body-fossils are absent. By con- trast, the known body-fossil record of oribatid mites com- mences during the Middle Devonian, but does not reappear until the Early Jurassic, at which time mite taxa are mod- em in aspect. All major plant taxa occurring in Pennsylvanian coal swamps, including lycopsids, spkenopsids, ferns, seed ferns and cordaites, were consumed by oribatid mites. Virtually every type of plant tissue was used by mites, notably indu- rated tissues such as bark, fibrovascular bundles and es- pecially wood, as well as softer seed megagametophytic and parenchymatic tissues within stems, roots and leaves. Sig- nificant evidence also exists for secondary consumption by mites of tissues in macroarthropod coprolites. Our data in- dicate that oribatid mites consumed dead, aerially-derived plant tissues at ground level, as well as root-penetrated tis- sues substantially within the peat. Oribatid mites were im- portant arthropod decomposers in Pennsylvanian coal swamps ofEuramerica. The wood boring functional-feeding-guild was expanded by insects into above-ground, live trees during the early Me- sozoic. New food resources for insect borers resulted from penetration of live tissues such as cambium and phloem, and the invasion ofheartwood and other hard tissues me- diated by insect-frngus symbioses. Termites and holome- taboUms insects were prominent contributors to this second wave of wood-boring, exploiting gymnosperms and angio- sperms as both detritivores and herbivores. An earlier em- Copyrrght ? 1997, SEPM (Society for Sedimentary Geology) placement of oribatid mites as detritivores of dead plant tis- sues continued to the present, but without a documented trace-fossil record. INTRODUCTION Oribatid mites are the most successful group of soil ar- thropods in temperate forest ecosystems (Johi?ton, 1982; Norton, 1994), where they are key animals?along with millipeds, isopods and coUembolan insects?responsible for the conversion of plant litter and wood to organic resi- dues in terrestrial decomposer food chains (Spencer, 1951; Harding imd Stuttard, 1974). Oribatid mites principally assist the decomposition of plant tissues by directly or in- directly interacting with the decomposer microbiota, es- pecially saprophytic fungi (Clement and Haq, 1984; Nor- ton, 1985; Afifi et al,, 1989), and thus assist in the commi- nution of plant and other biotal materials that eventually become integrated into the soil. The importance of oribatid mites is buttressed by their exceptional taxdc diversity worldwide and recurrii^ patterns of site-specific ecologi- cal dominance. Known oribatid mite diversity, excluding the Astigmata, consists of approximately 7000 nominal species within 1000 genera and 150 famihes, albeit the global tropical fauna remains largely uninvestigated (Nor- ton, 1990,1994). Although Early Devonian microarthropod mycophagy has been strongly indicated by Kevan et ai. (1975) and Sherwood-Pike and Gray (1985), there is sparse empirical evidence for microarthropods as major ^ents in decom- poser food webs during tiie Late Paleozoic (Rolfe, 1980). Notable exceptions to this general paucity of data include Banks and Colthart (1993) and Edwards et al. (1995), who discuss evidence for microarthropod-fungi-plant interac- tions from the trace-fossil record. There has been no com- prehensive community-wide synthesis of foss? evidence for the role of oribatid mites in the decomposition of plant material, with the exception of recent studies of Quater- nary environments, siunmarized in Ehas (1994). Current- ly, sufficient primary data exist that such a synthesis can be achieved for several, weU-collected, coai-swamp floras that were deposited in the Illinois Basin and the Southern 0883-13S1/97/0012-0319i$3.00 320 LABANDEIRA ET AL and Central Appalachian Basins during the Pennsylva- nian (Fig. 1). Whereas previous accounts of the presence of mite-bored woods and other tissues have suggested the oc- casional presence of oribatid mite decomposers (Baxen- dale, 1979; Cichan and Taylor, 1982; Stidd and Philhps, 1982; Scott and Taylor, 1983; Labandeira and Beall, 1990; Scott, 1977, 1992; Scott et al., 1992) esrtensive collections of perminerahzed coal-ball material now can supply a more comprehensive understanding of the extent and di- versity of plant-tissue degradation during narrow time in- tervals within the Pennsylvanian. There are four principal objectives of this report. First, we document new and extensive data indicating the per- vasiveness of oribatid mites as important agents of decom- position in Pennsylvanian coal-swamp forests of Euramer- ica. These data document an abundant and detailed trace- fossil record of postmortem tunneling into diverse plant tissues and taxa. Second, we evaluate the known body-fos- sil record of orifaatid mites, which we contrast with our new data on the fossil record of oribatid mite feeding trac- es. Third, we collate and summarize the existing hterature on the insect trace-fossil record of plant borings, and com- pare it to the corresponding oribatid trace-fossil record. Fi- nally, we place in spatiotemporal and paleoecological con- text the contrasting trace-fossil records of both oribatid mite and insect plant borers. From this comparison, we posit tentative conclusions regarding the taxic and ?colog- ie evolution of arthropod-mediated decomposition of indu- rated plant tissues in terrestrial ecosystems. ORIBATID MITES AND DECOMPOSITION Oribatid mites possess impressive ?cologie abundance, when gauged by the traditional measures of biomass and numbers of individuals. Most of the pioneering work and recent evaluations of the role that oribatid raites provide in the decomposition of various plant, fimgal, and to a less- er degree animal tissues, originates from studies of tem- perate to boreal ecosystems in the Holarctic and Palearctic realms. For example, oribatid mites are the overwhelm- ingly dominant macroscopic organism in central Europe- an rendzina soUs when evaluated as organismic biomass (Kubfl?ov? and Rusek, 1976) or in moder soils when cal- culated as fecal peUet biomass (Kubiena, 1955). In mea- sures of individuals per unit of volume or area, oribatid mites are the most abundant arthropod component (Sai- chuae et al., 1972; Wallwork, 1983; Eisenbeis and Wi- chard, 1987; Vasiliu and C?lug?r, 1987; Norton, 1990), at least for temperate forest communities. Wallwork (1957) determined from 2 to 15 mites/cm^ occurred in a vii^in hemlock/yellow birch forest in northern Michigan, where- as the same author later documented densities ranging from 5.6 mites/cm^ in mixed deciduous hardwood forest ft^m North Carolina, to 4.25 raites/cm^ in a Scots pine for- est from Sweden (Wallwork, 1983). Although minimally studied, oribatid mites are also abundant in the tropics. Lasebikan (1977, 1981) docu- mented three times more oribatid mite individuals than the next most abundant cohabiting arthropod group, ga- masine mites, in the terminal stages of a decaying palm trunk. This numeric abimdance is contrasted with depau- perate to modest taxic diversities at the few tropical sites that have been studied (e.g.. Beck, 1969,1972)--consider- ably less than the respectable 155 species in a plot of de- ciduous forest in the southern Appalachian Mountains (R. Hansen, pers. comm.). Within both tropical and temperate sites, the diversity of oribatid mites and other detritivo- rous arthropods is considerably less than that of superja- cent herbivore communities. This disparity is partly at- tributable to minimal resource partitioning by oribatid mites, which results from the greater importance of de- composition state than the taxic affinity of the food sub- strate (Forsslund, 1939; Riha, 1951; Hayes, 1963; Ander- son and Healy, 1972; Anderson, 1975, 1978). Neverthe- less, evidence indicates that some species possess quanti- fiable taxic-based specificity for particular food substrates (Hartenstein, 1962a; Berthet, 1964; Mignolet, 1971; Lux- ton, 1972). While species-rich clades commonly show only modest morphological diversity, across all taxa there is a substantial range of body form and mouthpart structure (Schuster, 1956; Norton, 1994), and most of the latter has yet to be functionally analyzed. Oribatid mites are small, chelicerate arthropods, and as adults range in size from 0.2 mm to 1.0 mm, rarely to 2.0 mm (Luxton, 1972; Wallwork, 1983), usually possessing a box-like exoskeleton superficially similar to that of a beetle. Individiials exhibit relatively long lives for mi- croarthropods, leisting ?:x)m one to four years for most spe- cies (Walker, 1964; Harding and Easton, 1984; Soma, 1990). Dietary correlates include poor nutritive value of ingested food, relatively low ingestion rates, and conse- quent minimal assimilation rates (Berthet, 1964; Luxton, 1972, 1979; Wallwork, 1983), which constrain their life- history parameters (Norton, 1994). These attributes are evidenced by their production of somewhat fewer than one to eight fecal pellets per day (Hartenstein, 1962b; Hayes, 1963; Berthet, 1964; Luxton, 1972). These pellets are inert objects that persist in the sou environment (Zachariae, 1965; Babel, 1975; Bal, 1982) and chemically differ mLni- mally from their parent plant tissues (Edwards, 1974). Or- ibatid mites form communities principally in decomposing plant htter and wood (Snow, 1958; Wooiley, 1960; Wall- work, 1976a) with some species acting as secondary con- sumers of dead but partially decayed plant material (Lux- ton, 1972; K?hnelt, 1976), others subsisting as copropha- ges on decomposer fecal pellets (Nicholson, et al., 1966; Wallwork, 1976b), and others, perhaps most, feeding prin- cipally or opportunistically on fongi (Forssland, 1939; Hartenstein, 1962a, b, c; Luxton, 1966, 1972; Shereef, 1971; Anderson, 1975; Andr? and Voegtlin, 1981; Wall- work, 1983; Clement and Haq, 1984; Norton, 1985; Afifi, et al., 1989). Most of this activity occurs in the L and upper F layers of temperate soils (the F^ layer of Zachariae [1965] and Bal [1968, 1982]) where accessible plant or- gans, such as conifer needles and dicot leaves, are con- sumed by endophytic and exophytic decomposers (Mitch- ell and Parkinson, 1976), leaving intact more resistant tis- sues such as needle epidermal sheaths and broadleaf vas- cular bundles (Zachariae, 1965; Babel, 1975; Bal, 1982). These substrates become degraded through a spatiotem- porai succession of ?cologie guilds dominated by oribatid mites, with each guild specializing on plant tissue of a par- ticular decomposition state (Harding and Stuttard, 1974; Wallwork, 1983). Oribatid mites mediate four major aspects of the degra- dation of plant tissues. First, they increase the surface of OniBATID MITES IN COAL-SWAMP FORESTS 321 plant tissues through fragmentation, digestion, and defe- cation (Kevan, 1962; Macfadyen, 1964). The resulting pel- lets are preferentially colonized by microbial decomposers, such as yeasts and bacteria, when compared to adjacent, undigested plant tissues. Second, the incorporation of de- composer fungi in fecal pellets allows for germination of some fungi within the fecal pellet substrate (Forssland, 1939; Shereef, 1971; Luxton, 1972; Ponge, 1984), often at some distance from the site of initial ingestion if the pel- lets sift through soil vugs or are biologically transported {Woodring and Cook, 1962; Wallwork, 1967; K?hnelt, 1976; Kilbertus and Vannier, 1979; Behan-Pelletier and Hill, 1983). The ecological significance of this role has been debated (e.g., see Woodring, 1965); Pande and Berthet, 1973). Third, oribatid mites create a "humus form" (M?U- er, 1879; Bal, 1973) in which fecal peUets within the soil become cohesive and fonn a pelletai matrix of larger sub- imits resistant to decomposition {Kubiena, 1955; Webb, 1977). This conglomeratic formation forms a distinctive stratal structure in many temperate soils and provides a biotope for further colonization by microarthropods (Ku- biena, 1955). Last, oribatid mites and other microarthro- pods and macroarthropods are pivotal in the vertical translocation of organic matter downward into the soil col- umn (Saichuae et al., 1972), partly as an effect of ?cologie guild succession during the decomposition process. Verti- cal translocation witi?n an "absorptively-saturated hu- mus" (K?hnelt, 1976:313) additionally enhances the effect of nutrient leaching by rainwater (Luxton, 1972). Many oribatid mites are foimd within decomposing structural material of plants (Seastedt, 1984; Ramani and Haq, 1991), and inhabit both hving and dead fungal fruit- ing bodies and hchens (Bellido, 1?79,1990). The specifici- ty of the association, and the exact nature in which the mites exploit the material, covers a broad spectrum. In particular, pieces of decaying wood usually support an abundant and diverse oribatid mite fauna (Trave, 1963; Swift, 1977; Seastedt, 1984). Most oribatid mites found in decaying wood are probably fui^vores (Norton, 1985; Seastedt et al., 1989), rather than strict xylophages, even though they may show moderate to high specificity for woody microhabitats {Wallwork, 1976a, 1976b). Along with other microarthropods, most extant oribatid mites hving in wood or other indurated plant tissues inhabit in- terstices created by biotic factors {K?hnelt, 1976), such as space created by larger tunneling organisms or the wedg- ing of penetrating plant roots. Of those mites that con- sume wood, many exploit these ambient spaces rather than tunnel {Trave and Duran, 1971). Oribatid mites that create hving spaces by their feeding activity may do so in both adult and immature instars or, more commonly, only as immatures. They frequently cre- ate irregular feeding galleries that can become spacious relative to the size of the mite. Alternatively, true tunnel- ing may occur, in which the living space is an elongated tube, gradually increasing in diameter as the mite grows {Wallwork, 1957), and packed with fecal pellets behind the active feeding region (Jacot, 1936; Gourbi?re et al., 1985; Lions and Gourbi?re, 1988,1989), For some lineages, such as the Lohmanniidae (Haq and Konikkara, 1988), the type of endophagy is intermediate between these types, with tunnel-Uke areas produced by communal feeding. In gen- eral, oribatid mites tend to have broader microhabitat ranges and possess more eclectic feeding habits as adults than as immatures. The diverse feeding patterns of oribatid mites have been characterized in three major ways. Initially, Riha (1951) considered oribatid mites as generahsts or speciahsts. Schuster (1956) later categorized the more specialized or- ibatid mites as microphytophages subsisting on microbi- otal elements, and macrophjrtophages that consume high- er plant tissues. K?hnelt (1976) characterized oribatid mites as endophages, subsisting in internal plant tissues or ectophages, as external feeders. Schustei^s widely ac- cepted categorization, was expanded by Luxton (1972) to include the more host-specific coprophages, zoophages, and necrophages, feeding respectively on fecal material, live animals, and dead animals. Zoophages and necropha- ges seem relatively inconsequential as oribatid mite di- etary classes (Wallwork, 1983; Norton, 1990); coprophagy apparently occurs with greater frequency hut is poorly documented (Schuster, 1956; Anderson and Healey, 1972; Luxton, 1972; but see Wallwork, 1967,1976a). Macrophy- tophagy principally includes leaf and needle fitter con- sumers, and wood consumers. Although cellulose is digest- ed by various oribatid mite species with either intrinsical- ly produced (Wallwork, 1983) or more Hkely microhiafiy- borrowed celluiytic enzymes (Ziakler, 1971; Stefaniak and Seniczak, 1976, 1981; Norton, 1985, 1994), lignin appar- ently is not digestible (Woolley, 1960; Harding and Stut- tard, 1974). Those macrophytopha^es feeding on wood ini- tiafiy probe moist and fiingally prepared wood by entering clefts and fissures (Trave, 1963; K?hnelt, 1976) and com- mence boring at the teimini of these cracks, avoiding resin canals during timneling (Jacot, 1936), and in some cases entering heartwood (Pande and Berthet, 1973; Wallwork, 1976b). Recent experimental work (Scheu and Schulz, 1994; Schulz and Scheu, 1994) indicates that the term phytosaprophagy may be the most accurate description re- garding the effect of mites on wood decomposition (see Woas et al., 1989). Macrophjrtoph^es feeding on foliar material preferentially devour parenchymatic tissues. Co- nifer needle fragments and petioles of dicot leaves are transformed into sacs of fecal pellets and imdigested vas- cular tissue that are surrounded by epidermal sheaths (Gourbi?re et al., 1985,1987; Lions and Gourbi?re, 1988, 1989), which eventually rupture as they are transported deeper into the sou horizon (Zachariae, 1965). Dicot leaf blades are skeletonized exophytically by attack through the leaf cuticle, resulting ia the casting of fecal pellets on the substrate surface (Riba, 1951). Although our com- ments have emphasized that oribatid mites overwhelm- ingly consume fungi or dead plants, recent evidence indi- cates that many species are facultative herbivores (Wall- work, 1965, 1967; Ramani and Haq, 1984; Walter et al., 1994), mining the leaves {Coulson, 1971; Norton, 1983; Karg, 1984; Fernandez and Athias-Binche, 1986) or boring the twigs (Lan et al,, 1986; Walter et al,, 1994) or roots (Ja- cot, 1934; GofFand Blanchi, 1983) of live angiosperms, FOSSIL HISTORY OF ORIBATID MITES Several lines of evidence attest to the geological antiq- uity of oribatid mites, including Pangaean biogeographical distributions of many extant genera (Hammer and Wall- work, 1979; Wallwork, 1979), a mosaic pattern of mor- 322 LABANDEIRA ET AL Perio?or Ma Era Subperiod I Body Fossils Or?batid Fossil Record I Trace Fossils of Insect Wood Borers Trace Fossils o o > N n O *^ c m 50- O H in 3 O lU 100- o nt 0) ^ O ? O N 150- O Q> u S 0) 200- O w 0> ? ^ 1- 250- c n ? c 300- u d> 4DD- a 17, h?30. Pleistocene: conifer, Cole?ptera I 31 - Mes&iJiJan: dicot, DipteJ'a [7^32. ToflDnian: dicot, Coteoptera (Ceraniljycidae) ?33. Sermvalianr monocot, C?l^apler? ?34. Langhian: ?. Cole?ptera <^5. Aquhtanian: coniler. Hymenoptera {Siricjdae) ?^^^6. Chattian: conifer, EsopTera ^Kalotermitidae) ^^^37^ ?LrpgIJan: con?er. Cole?ptera ????3fl- Bartonian: dtcot. Cole?ptera SJ??'39- Eocene: dic?tn D?ptera {Agrom^Hae) 10 11. la. 40r Than?tian? dtcot. Hyrrwnoptsra (FormicldBe) 41. MaaGtrJctian: dicotn Isoptera 42. Cenomanian: conifer. Isoptera 43. BarTfimfan: conilsr, CoJeoplera (ScoF^i?ae) 44. Ki?Ti?reridgian: pleridosperm?, CQI? ?ptera (Cupedidae) 45. B^oc?an; coniler. Cole?ptera? 4&. Lies: cdnrfer. Cole?ptera (Anobi?dae) 47. Carnian^ coniler. Cole?ptera? 4S. Carnian; contter. Cole?ptera Midcontinent and Appalachian Coal- Swamp Floras {Fig. 2) 2t. Caihoun 22. Henri n ? 23. Rockspring 24, Upper Path Fork 25. Newcastle FIGURE 1?The documented fossil records of oribatid mites, and on batid mite and Insect wood-borings. Three panels of fossil occurrences display, from left to fight, the oribatid mite body-fossil record, the trace-fossil record of oribatid mite borings, and the Insect trace-fossit record of borings. Note the occurrence of Cattoniferous and Early Pemiian oribatid mite borings to the exclusion of the fossil records of either oribatid mite body fcsslis or of insect borings. Solid bars represent secure identifications and stippled bars represent probable Identifications; question marks indicate some geochronologic uncertainty in stage placement within an epoch. The geochronology is from Harland et al. {1990). First panel {literature sources 1 to 16) represents the body-fossil record of oribatid mites {Krlvolutsl^ and Druk, 1986; Krivolutsky et al., 1990). See Appendix 1 for details. Data sources are: l-Quatemary; Hopkins et al., (1976), Schelvis (1987, 1990), and Erickson (1988); 2-Ule Pliocene (Piacenzian); Gosolova et al., (1985); 3-Early Pliocene (Zanclian); Gosolova et al., 1985); 4-IVIIddte l^iocene {Serra val lian); Pampaloni {1902); 5-Early l^/ltocene {Aquttanlan); Woolley (1971); S-Late O?gocene (Chattian); Peinar {1992); Norton and Poinar. (1993); 7-3pproxtmateiy Early Oligoc?ne (Rupellan); Seilnick (1919, 1931); 8-Late Paleoceno {Thanetlan); Baker and Wighton (1984); 9-Early Paleoceno (Danian); Krivolutsky and Ryabinln (1976); 10-Late Cretaceous (Campsnian); lulcAipine and Martin (1969); 11-Late Cretaceous (Santonian); Bulanova- ORIBATID MITES IN COAL-SWAMP FORESTS 323 phology characterizing primitive taxa (Krivolutsky et al., 1990; Woas, 1991), and notably their fossil record {Sell- nick, 1927; Seiden, 1988, 1993b; Dubinin 1991), particu- larly the presence of early derivative taxa from the Middle Devonian (Norton et al., 1988, 1989) and extant genera known from deposits as old as the Jurassic (Sivhed and Wallwork, 1978; Krivolutsky and Ryabinin, 1976; Nied- baia, 1983; Krivolutsky and Druk, 1986). Although oriba- tid mites, as a clade, were present during the Early Devo- nian, one particular and important subclade, the Astig- mata, originated within the oribatid lineage some time lat- er, probably during the Late Paleozoic (0 Connor 1982, 1984), Astigmatic mites currently comprise about half of the species-level diversity of oribatid mites (Norton, 1994), and are consummate exploiters of plants, insects, and ver- tebrates. Of all arachnid groups, the fossU record of non- astigmatic oribatid mites is unique both in terms of the relatively good representation of adult instars with hard- ened exoskeletons (Bernini, 1986) and the occurrence of characteristic borings in indurated plant tissues, especial- ly wood (Scott and Taylor, 1983). Ironically these two as- pects of the oribatid mite fossil record?the body-foss? rec- ord of adult morphology and the trace-fossil record of be- havior and life-habits?are geochronologically comple- mentary (Fig. 1). Whereas the earhest known body fossils originate from a Middle Devonian deposit, the body-fossil record does not resume until after a hiatus of 170 million years, during the Early Jurassic, after which occurrences continue sporadically to the Recent. By contrast, the trace- fossil record is richest during much of that 170 miUion year gap, especially dmring the Carboniferous. Thus, the oribatid mite fossil record, although sparse and woefully understudied, offers a coarse chronology of both very early and essentially modem lineages (Krivolutsky, 1973), as well as a glimpse into the plant-feeding ecology and behav- ior of taxonomically unknown Paleozoic forms. The Body-Fossil Record Fossils of oribatid mites occur in three preservational modes: as compact and unaltered cuticle, permineraliza- tion in amber, and less commonly as compressions or im- pressions in mudstones. Both the oldest known oribatid mite fossils, described from Middle Devonian strata, and some of the youngest mite fossils, found as Quaternary subfossil material, are preserved as original cuticle. The oldest fossil material is highly flattened and exhibits somewhat disarticulated sclerites, retainingmost append- ages (Norton et al., 1988, 1989; see also Kethley et al., 1989), whereas subfossil material is three-dimensionally preserved but often with missing major appendages (Eli- as, 1994). These two differing results are attributable principally to raicrofossil processing techniques: Devonian rock is macerated in hydrofluoric acid and successively ex- posed layer by layer, whereas Quaternary sediment is typ- ically sieved, residting in a loss of appendages from abra- sion. By contrast, preservation of oribatid mites in amber retains entire three-dimensional bodies, even of poorly sclerotized lineages and subadult instars (Krivolutsky and Druk, 1986). However, oribatid mites preserved in amber are known only from Late Cretaceous to Pleistocene ma- terial (Womersley, 1956; McAlpine and Martin, 1969; Aoki, 1974; Schl?ter, 1978; Larsson, 1978; Poinar, 1992). Rare examples of oribatid mites are known from compres- sions and impressions in mudstones (Baker and Wighton, 1984; P?rez, 1988). In total, 104 pre-Pleistocene oribatid mite fossils have been described (Appendix 1), although many times this number have been recorded in Quater- nary deposits (Elias, 1994). Quaternary oribatid mite pa- leoecology is a vast discipline that emphasizes use of fos- sils of modem species in establishing the recolonization dynamics of deglaciated Pleistocene landscapes (Karpin- nen and Koponen, 1973; Elias, 1994), seasonal variation in paleolimnological studies (Erickson, 1988), and evaluation of site-specific conditions at Holocene archaeological sites (Schelvis, 1987,1990). The earliest known oribatid mite fossils are Pro- tochthonius gilboa Norton and Deuonacarus sellnkki Nor- ton from the Middle Devonian Gilboa mudstones in south- central New York state (Fig 1; Appendix 1; Norton et al., 1988,1989). Each of these species comprises a monobasic family that is primitive in many respects, yet still exhibits Zachvatina (1974); 12-Lat6 Cretaceous {Turanian); Grima?di, pers, comm.; 13-Early Cretaceous (Albian); Schlee (1972); Grimaldi, pers. oomnn.; 14-Late Jurassic (Tithonian); Krivolutsky and Ryabinin (1976); 15-Early Jurassic (Sinemurian); Sivtied and Waliworit (1978); and IB-Middle Devonian (Givettian); Norton el al. (1988, 1939). Second panel (literature sources 17 to 29) represents the trace-fossil record of oribatid mite borings. Data sources are: 17-Qijatemary; Haari?v (1967) and Koponen and Nuorteva (1973); 18-Earty Cretaceous (probably Barremian); Seward (1923, 1924); 19-Middte Jurassic (Aalenian); Vao et al., (1991); 20-Eariy Pemiian {stage not designated); Goth and Wilde (1992); 21-Calhoun Coal-Ball Flora, early Late Penn- sylvanian (Chamovnicheskian); this report; 22-Hemn Coal-Ball Flora, late Middle Pennsylvanian (Myachkovskian); this report; 23-Rock Spring Coal-Ball Flora, middle Middle Pennsylvanian (Podolskian); this report; 24-Upper Path Fork Coal-Ball Flora, early Middle Pennsyivanian {Ka- shirskian); this report; 25-New Castle Coal-Ball Flora, Early Pennsylvanian (Cheremshanskian); this report; 26-Early Pennsylvanian {Kinder- scoutian); Labandeira et al. (1994); 27-Middle Mississippian (Asbian); Chaloner, et al., (1991); 2B-Middle MIssissippian (Asbian); Rex (1986), Rex and Gaitier (1986); 29-Late Devonian (Famennian); Arnold (1952), F.M. Hueber and Labandeira, (pers, otssen/.). Third panel (literature sources 30 to 48) represents tne trace-fossil record of insect borings. Family designations in parentheses designate the earliest, well-established record as trace fossils. Additional Cenozoic occurrences have been omitted for lack of space. Data sources are: 30-Quaternary; Scudder (1900); 31-Late Miocene (Messinian); Suss and MQIIer-Stoll (1975); 32-Late Miocene (Tortonian); M?ller-Stoll(1989); 33-Midd!e Miocene (Serravatlian); Call and Tidwell (1988); 34-Middle Miocene (Langhian); Guo (1991); 35-Early Miocene (Aquitanian); Roselt and Feustel (1960); 36-Late Oligoc?ne (Chattian); Rozefeids and Oe Saar (1991); 37-Early Oligoc?ne (Rupelian); Bachofen-Echt (1949), Larsson (1978), and Krzemi?ska et al. (1992); 38-Midd)e Eocene (Bartonian); Gregory (1969); 39-Eocene (stage not designated); Suss and M?l?er-Stoll (1980); 40-Late Paleocene (Thanetian); Brues (1936); 41-Late Cretaceous ?Maastrichlian); Rohr, et al. (1986); 42-Late Cretaceous (Cenomanian); Labandeira (pers. obsen/.); 43-Early Cretaceous {Banemian); Blair (1943) and Jarzembowski (1990); 44-Late Jurassic (Kin:?- meridgian); Tidwell and Ash (1990); 45-Middle Jurassic (Bajocian); Zhou and Zhang (1989); 46-Lower Jurassic (stage not designated); Jurasky (1932); 47-Late Triassic (Carnian); Walker (1938); 48-Late Triassic (Carnian); Linck (1949). 324 LABANDEIRA ET AL derived features. The known body-fossil record then paus- es for 160 million years, and resumes during the Early Ju- rassic, in which two taxa referred to modem genera were described by Sihved and WaUwork (1978). The modem morphology of these Jurassic specimens (Norton, 1990) provide a stark contrast to Middle Devonian taxa that are assigned to extinct families. Body fossils of modem fami- lies also have been described from Late Jurassic, Late Cre- taceous, and Cenozoic deposits {McAlpine and Martin, 1969; WooUey, 1971; Bulanova-Zachvatitia, 1974; Krivo- lutsky and Ryabinin, 1976; Krivolutsky et al., 1990; Nor- ton and Poinar, 1993). Additionally, of the 70 oribatid mite species described from Late Eocene Baltic amber (SeU- nick, 1919,1931; Keilbach, 1982), nearly all are assignable to modem genera (Krivolutsky et al., 1990). This pattern of taxic longevity again suggests that oribatid mites are an ancient group. The Trace-Fossil Record The abundance of characteristic Mississippian to Early Permian borings in plants and coprohtes indicates that or- ibatid mites exhibited significant levels of tunneling be- havior in diverse vascular plants. These well-preserved permineralized trace-fossils provide unique insights into oribatid mite microhabitats, life-styles, and plant host preferences that are unavailable for any other part of the foss? record. The earhest knoviTi borings that are conceiv- ably attributable to oribatid mites are undescribed tun- nels occurring in Prototaxites, an enigmatic terrestrial or- ganism, probably a basidiomycete (F. Hueber, pers. comm.), of Late Devonian age from New York state (Ar- nold, 1952). These Prototaxites "logs" contain simple, line- ar to undulating, occasionally joining tunnels in indurated mycelial tissue (F. Hueber and CCL, pers. observ.) that ex- hibit evidence of a reaction rim on the tunnel inner smface and contain coprolites of digested mycelia within the tun- nels. However, tunnel diameters are several times wider than modem analogs, such as carabodid mites, which typ- ically bore into living and freshly dead basidiocarps (Mi- chael, 1882; Reeves, 1991, 1992) or into hchenized fungi (Bellido, 1979; Gjelstrup, 1979; Gjelstmp and S0chtig, 1979), producing dense networks of tunnels. From Car- boniferous and Lower Permian deposits, suspected and well-confirmed oribatid mite tunnels and coprolites have been documented from every major plant group, including lycopsids (Scott, 1977; Chaloner et al., 1991), calamites (Taylor and Taylor, 1992), ferns (Lesnikowska, 1989; Scott et al., 1985,1992; Rex and Galtier, 1986), seed ferns (Stidd and Phillips, 1982), cordaites {Cichan and Taylor, 1982; Scott and Taylor, 1983; Rolfe, 1985; Labandeira and Beall, 1990), and conifers (Goth and W?de, 1992). PubHshed ac- counts occur from at least 10 separate horizons spanning a 75-niillion-yeaT interval, mostly ?x>m clastic- and peat- dominated swamp forests of Euramerica (Fig. 1). Al- though previous reports were largely anecdotal, in this re- port we expand documentation of both the intensity and diversity of tissues that were degraded by these early de- composers. While exquisitely permineralized tissues of Pennsylva- nian swamp forests provide an important, 17-niillion-year record of the early history of wood-borers, this taphonomic window apparently closed duribag the Early Permian (Fig. 1). Subsequently, evidence for wood-borers becomes avail- able for the Late Triassic (Walker, 1938; Linck, 1949), and the frequency of relevant fossil deposits increases toward the present. However, the Late Triassic to Recent record of terrestrial wood borers is exclusively insect-driven, con- sisting overwhelmingly of Cole?ptera and Isoptera, with subordinate occurrences of Hymenoptera and Diptera. These two, chronologically nonoverlapping records of tax- onomicaliy distinct arthropods undoubtedly represents the effects of differing taphonomic regimes in the fossil record, and in particular, large accumulations of degrad- ed, often woody plant tissue during the Carboniferous, when compared to the post-Paleozoic (Robinson, 1990; La- bandeira et al., 1994). Nevertheless, the emergence of new Uneages of insect wood-borers during the Triassic may represent a profound shift ftx)m Late Paleozoic oribatid mites to Mesozoic and Cenozoic insects as the principal de- tritivores of Hgnified, above-ground plant tissue. Signifi- cantly, some of these insect Uneages were dietarily herbi- vores, consuming live tissues adjacent to wood, such as phloem and various meristems. Oribatid mites are the only arachnid group with both re- spectable body-fossil (Appendix 1) and trace-foss? records (Figs. 1-12). Yet these dual fossil records are separate and have not been linked, either by instances of tunnels har- boring mite body fossils or by mite body fossils containing coprolites with identifiable plant tissues. It is possible that these independent fossil records eventually may he linked, although the relative preservation potential of coexisting plant and arthropod cuticle is poorly understood. MATERIALS AND METHODS Bulk tissues from plants occurring in peat-dominated coal-swamp forests of equatorial Eiiramerica were fre- quently permineralized in sedimentary basins occurring at or slightly above sea level. These carbonate-perminer- alized tissues characteristically occur as coal ball zones in relatively planar coal deposits of Pennsylvanian age. The Pennsylvanian deposits examined in this study corre- spond to an absolute age from 315 to 298 million years (Fig. 2; Harland et al., 1990), representing a 17 Ma inter- val. Preservation of these plant tissues reveal anatomical detail at the whole organ, tissue, and individual ceUular levels. While coal balls can range in maximum dimension ftxim several centimeters to more than 1 meter in the field, vir- tually all coal balls used in this study were at the small end of this range. Individual coal balls were rock sawed in the laboratory into slabs that were from 2 to 6 cm thick. Surfaces of these slabs were smoothed by grinding in moistened 500 grit carborundima powder, followed by acid etching in 5 percent hydrochloric acid for 12 to 17 seconds (for details, see Phil?ps et al., 1976). Acid etching exposed organic material on the coal-ball surface, such as ceU walls, as it removed carbonate matrix. After the etched surface was afr-dried, the surface was flooded with ace- tone, followed Lmmediately by roiling a 2 or 3 mil sheet of transparent cellulose acetate onto the surface. Once the sheet was dry, it was ?fted off, taking with it the embed- ded, thin layer of organic material exposed on the surface. This procedure was repeated until a biologically meaning- ful thickness of the coal-ball surface was captured as peels. ORIBATID MITES IN COAL-SWAMP FORESTS 325 M?C,^-. ^?^?:?|^!^:1^f??t^g^^:'?;?#?t'-^:;l^;?^r? - Vi^^?-"' -?^^ ' :^i^iry''.'^f ? ." i L 1 :v; z 1 Geochronolnrjy 1 Hartandstal., 1990 ? Illinois 1 Appalachian Pennsylvan. 1 Successions 1 USA fUFQpir 1 1 1 .1 Basin ^? Basins to 1 1 CO is ? 1 CO ? to (D 05 p 3 < ? z G Z O :? tu Aodaiaehian j-r-A?L\-j-i.^-^^^ / 1 2 CQ u 7 en o o ED ? o o CO O E 1 m y i y ^ Upper Path Fork Coal ? z < i m cr^. i MO W i . 1 ' ' I- :?: >- m IJ ! ? 1 . llllnot? Vi?] Bas=n- H-i- sJi j z z tu '1.. 310- c CO lU S?^^ /?il 1 1 tu O 5 f V ? ' 'J' " ?r\^ 1 1 Nv'-'^^-t^?'^^?J^ 1 1 '^. (L G3?1 ?: T-L I i-ip 1 ? m ^-^ __^ _l ? ' UJ Ul ?R J /? ^ > > o ^1? ?r?- 1 EL ? < .:J^. "^ td O lu ' IT 2 3 i? u < CD Z Z Z Ul }iir c ? CL ? Ul CTt 7 .Ti" C ? ?m ?'^ ?D U < Z 'i ti ....;?.,.? .. p;^- ?-f.^ii ly it?_ -- ., .-? ^A:^4:???;j-t ;^.-i:.i,-. .. ;;..;???-: V.. ? ? ? - ^--^ -, ?.?.--.-.^??.^y?,?-,.._.- ?: -^ ^^ ?^^^ ?" . :?,-.?.. ?. .--^ ."..' .. ?- .? si.ix.'^''-^^ . ? -:-\ :?:-*-, -y -.i' '??> .:-?- . ..\ FIGURE 2?Stratigraphie, geochronologic, and paleogeograptiic context of coal-ball deposits referred to in text. Geochronotogic calibration is Irom Harlancj et al. (1990); positions of Pennsylvanian basins are from Cross and Phillips (1990) and Winston and Phillips (1991 ). Abbreviations: Garb. = Cart>ondate Formation, Cass. = Casselman Formation. For light microscope photography, regions of interest firom the acetate peels were cut and moimted in embedding me- dium on microscope slides. An Olympus SZH microscope connected to an automatic photographic system was used to photograph slide-mounted peels. Three dimensional reconstructions of mite borings in several types of plant tissues were based on camera lucida drawings of cellulose acetate peels. Initially, a slab with well-preserved plant tissues containing mite borings was selected, and then from 100 to 150 successive peels were 326 LABANDEIRA ET AL FIGURE 3?Three-dimensional reconstruction of mite galleries in leal cushions of the lycopsid LepidopNolos har?ourt?i imm the Central Ap- palachian Basin. Specimen 28969-B bot. Stippled pattern represents leaf cushion parenchyma; elliptical objects in black represent pa rich nos. Mite e^ccavated areas in leaf cushions were rendered without coprolites and tissue fragnnents for ease of viewing. Reconstmctions were based on peel slices 1 to 24 (a), peel slices 25 to 49 (b), peel slices 50 to 74 (c), and peel slices 75 to 99 (d). Light photographs o? peels No. 1 and 25 are attached to (a) and (b) respectively. made. From each series of peels originating from a slab surface, every tenth peel was selected as a control. A cam- era lucida line-drawing of each control peel was made at 6 X magnification on tracing paper, with plant tissues and mite-assodated features labeled. Each successive camera lucida Une drawing then was assembled, integrated, and three-dimensionally rendered by use of Spyglass Dicer" and Adobe Illustrator' software on a Macintosh Quadra computer. After the resulting printout was manually re- produced on mylar, it was photographically reduced for in- corporation as elements in Figs. 3 and 4. All coal ball specimens, acetate peels, and associated mi- croscope slides reside at the Paleobotanical Research Cen- ter of the University of lUinois, Champaign-Urbana. Peels that were used for supplying microscope sUdes in this re- port are the following: 28938-Btop 28969-Bbot 30679-Bbot 31024-Btop 31272-Bbot 31272-Ctop 32274-A3-Side2 32350-Cbot 32479-A 32479-Bbot 32479-Ctop 37112-Etop 37112-Itop 37326-Gbot 38040-Gbot 38043-Lbot 38066-Ctop 38070-Etop 38078-Ctop 38374-Btop 38600-Bbot Microscope shdes associated with the above peels that 10765-Cbot 32274^15A-4top 38012-Dbot following; 28686-A 32277-Dtop 38022-Etop 22,329 22,438 22,577 28799-Btop 32330-Bbot 38030-Ktop 22,354 22,441 22,608 28892-Btop 32330-Btop 38035-Hbot 22,363 22,446 22,612 ORIBATID MITES IN COAL-SWAMP FORESTS 327 FIGURE 4?Mfte tunnels in Recent plant tissues (a-e), republished with permission of the authors, (a) A twig of Quercus (oak) from the A hoiizon of a moder rendzina soil, showing consumption of wood. From Kub?kov? and Rusek (1976, Plate 13, figure 1; X15). (b) An unidentified fragment from a calcareous marl in a rendzina soil from Sierra Grossa, Spain. From Kubiena (1955, figure 13; x 30). (c) Deutonymph of Hoptophoretla thoreaui (Jacot) in a Picea (spruce) needle. From Jacot (1939, figure 1; length of mite about 0,5 mm), (d) A needle of Pinus (pine) containing a tunnel with fecal pellets. From Ponge (1988, figure 1; scale bar = 1 mm), (e) Fragment of Pinus wood from a litteryhumus horizon, tunneled by Rhysotritia duplicata. From Dhft (1964, figure 15; xia). Middle Pennsylvanian wood (f-g) with mite galleries, (f) A fragment of calamitean wood, from the Rock Spring Coal-Ball Flora. Peel No. 9 o( coal ball 3233-Btop (slide 22,407), Scale bar = 1 mm. (g) Three-dimensional rendering of tunnel network for peels 1-120 in f. 328 LABANDEIRA ETAL 22,393 22,456 22,643 22,400 22,471 22,662 22,403 22,492 22,667 22,406 22,515 22,670 22,407 22,517 22,672 22,416 22,520 22,673 22,428 22,524 22,727 22,433 22,537 Locality and stratigraphie infonnation for the seven coal-ball floras investigated were based on superpositional relationships provided by Ph?lips (1980) and PMUips et al. (1985), and temporally calibrated by Harlan et al. (1990). The examined coal-ball floras are as follows, from oldest to youngest. The New Cas?e Coal-Ball Flora is located in the Southern Appalachian Basin, near Townley, Walker Co., Alabama. It is of Cheremshanskian age (about 315 Ma) within the Early Pennsylvanian Subperiod, and is assign- able to the Westphalian A stage of classic European no- menclature. The Upper Path Fork Coal-Ball Flora is locat- ed in the Central Appalachian Basin, from the Cranks Creek locality, near Cawood, Har?an Co., Kentucky. It is of Kashirskian age (about 308 Ma) within the Middle Penn- sylvanian Subperiod, and corresponds to the Westphalian B stage of classic European terminology. The Rock Spring Coal-Ball Flora is located in the Central Appalachian Ba- sin, from the Cross Mountain locahty, near Careyville, Campbell Co., Tennessee. It is of Podolskian age {about 306 Ma) within the Middle Pennsylvanian Subperiod, and corresponds to the Westphahan C stage of classic Europe- an temunology. The Murphysboro Coal-BaH Flora is locat- ed in the Illinois Basin, from the Maple Grove strip mine, near Cayuga, VenniUion Co., Indiana. It is of Podskohan age (about 305 Ma) within the Middle Pennsylvanian Sub- period, and corresponds to the early Westphalian D stage of classic European terminology. The Herrin Coal-Ball Flora is located in the Illinois Basin, from the Carrier M?ls (Saline Co.) and Shawneetown (Gallatin Co.) locali- ties, nUnois. It is of late Myachkovskian age (about 303 Ma) within the Middle Pennsylvanian Subperiod, corre- sponding to the late Westphalian D stage of classic Euro- pean temunology. The Baker Coal-Ball Flora is located in the Illinois Basin, from the Hart and Hart Coal Company Mine, near Providence, Webster Co., Kentucky. It is of late Myachkovskian age (about 303 Ma) within the Middle Pennsylvanian Subperiod, and corresponds to the late Westphahan D stage of classic European terminology. The Calhoun Coal-Ball Flora is located in the Dlinois Basin, from the Calhoun and Benyville localities, Kichland Co., Illinois. It is of Chamovnicheskian age {about 298 Ma) within the Late Pennsylvanian Subperiod, and corre- sponds to the Stephanian A stage of classic European ter- minology. Additional details regarding ioca?ties and tem- poral occurrences of these floras are provided in Phillips (1980) and Fig. 2. RESULTS Plant tissues from seven important coal-ball floras from Pennsylvanian deposits of equatorial Euramericawere ex- amined for damage by borers. From these seven deposits (Tables 1 imd 2A; Figs. 3-12) the foUowing were docu- mented: (i) the size and shape of coproHtes occurring in tunneled and galleried tissues, (ii) the incidence of tissue- boring, (iii) identities and percentage of the major plant taxa consumed by detritivorous borers, (iv) the spectrum of plant organs and tissue types preferred by tissue-borers for each coal-ball flora, and (v) the degradational condition of plant tissues at the time of consumption. These features are presented below as a temporal overview of the taxic, histologie, and edaphic patterns of borer-mediated plant damage among the earhest, well preserved, coal-swamp floras. The coprolites documented in our study bear smooth surfaces and range from 47.3 to 106.6 ^i.m in diameter {Ta- ble 2A). Thefr diameters are within the size range report- ed from Middle Mississippian (Esnost and Roanne, France) to Lower Permian (Wetterau, Germany) permi- neralized floras of several previous studies (Table 2B), some of which have attributed thefr coprolite producers to oribatid mites (Baxendale, 1979; Scott and Taylor, 1983; Goth and Wilde, 1992). Additionally, clusters of coprohtes in many plant tissues often possess a stereotyped shape, with individual groupings having ovoidal, eUiptical or rarely cylindrical shapes. Diameter-to-length ratios vary from approximately 1,6 for ovoidal shapes to approximate- ly 2.8 for cylindrical shapes. These coprohtes are always found in borings ranging in diameter from 100 to 450 jjon, and within dead plant tissues that include diverse types of parenchsrma, phloem and xylem, and once-digested plant material of other detritivores and herbivores. Coprolites of the 40 to 110 \i.m size class are almost always associated vnth endopbytic detritivory (Table 2). In Table 1, we determine the total percentage of peels with coprohte-bearing tunnels or galleries for all seven coal-ball floras. We list also the percentage of peels con- taining borings in tissue fragments from the five major plant taxa: lycopsids, sphenopsids, ferns, seed ferns, and cordaites. To determine the incidence of detritivory for each coal-ball flora we tabulate the number of identifiable tissue fragments from major plant taxa that contain de- monstrable evidence of mite boring, and standardize these counts across aE major taxa for each coal-ball flora. Thus each peel, which lacks or contains single or multiple mite- bored tissue fragments, contributes to the frequency data presented in Table 1. Comparison of previously gathered biomass abxmdance of major plant taxa for six of ?iese flo- ras {Phillips, et al., 1985; Winston and PhiUips, 1991) with the corresponding incidence of borer damage for each plant group provides an assessment of whether borer de- tritivores are temporally tracking taxic trends in plant biomass. The earliest well-documented coal-swamp deposit from North America is the New Castle Coal-Ball Flora, of Early Pennsylvanian age {316 Ma) from northwestern Alabama, in the Southern Appalachian Basin. Plant biomass from this deposit is overwhelmingly dominated by lycopsids (82.2 percent), and the subordinate volumes of sphenop- sids, fems, and seed ferns collectively represent 5.0 per- ORIBATID MITES IN COAL-SWAMP FORESTS 329 TABLE 1?Incidence ol mite detritivory from North American coal-swamp lloras of Pennsylvania Age. Num- ber Perce- Num- of peels with ntage of peels Total num- Coal- Ball Coal-Ball peels ber of peels tun- nels & with ber tun- of neis & feed-. Total biomass percentage contribution of major plant taxa^ and extent of detritivory^ for each major taxon (in parentheses) Shoot Coal-ball flora collec- tion' exam- ined^ exam- ined COpro-copro- lites lites ing traces Lyocpods Sphenop- sids Ferns Seed ferns Cordaites to root ratio' Calhoun VSl all 370 106 Baker Herrin VS5 RS2 ES middle middle middle 518 272 93 116 100 71 25.0 36.8 292 139 1.9 (5.5) {52.5) 9.3 (0.3) (8.6) 64.3 (76.7) (7.2) 24.2 (15.1) (15.1) 0.3 (2.4) (16.5) 0.6 Miiiphysboro Rock Spring Upper Path Fork VS4 VS9 RS 1 RS 1 middle middle all all 374 1360 58 33 47 606 10 22.3 18.2 44.6 163 42 833 68.9 {52.8) 76.0 {73.8) 50.9 (21.8) 4.1 (3.7) 3.0 (0) 9.4 (17.9) 10.1 (14.7) 6.5 (4.8) 4.0 (1.9) 16.3 (10.4) 1.0 (0) 2.7 (2.4) 0.5 (16.6) 14.5 (16.7) 33.0 (55.9) 1.89 1.69 0.72 New Castle RS2 RSI middle all 301 408 58 74 18.9 18.1 70 72 54.5 (62.9) 82.2 (95.8) 3.6 (12.9) 1.2 (2.8) 3.1 (12.9) 1.0 (1.4) 2.1 (4,3) 2.8 (0) 36.7 (7.1) 0(0) 0.47 1.8 ' Abbreviations: VS, Vertical Section; RS, Random Sample; ES, Ecology Studies. ^ A middle peel represents a surface located approximately in the central region of a coal ball. For each coal ball, a middle peel was selected from a series of serveral peels from successively sawn coal-hall slices. ^ Floristic data frx>m Phillips, et al. (1985) and Winston & Phillips {1991). Italicized, entries represent floristically dominant taxa by biomass. Floristic data was not collected for the Baker Coal. * Data from frequency counts of specimens with coprolites in borings, normalized to 100 percent. Detritivory in macrocoproUtes and unknwon tissues omitted. ^ The shoot-to-root ratio ia the proportion of biomass occurring as above-ground tissues versus below-ground tissues in the coal-ball ?oras examined. cent of the deposit CWinston and Phillips, 1991). Cordaites are absent. Although the percentage of peels with evi- dence of mite detritivory across all taxa was 18.1 percent, lycopsids were overwhehningly attacked by detritivores, accounting for 95.8 percent of aU plant tissue fragments with mite tunnels. The principal plant hosts subjected to borer attack was the lycopsid Lepidophloios cf. harcourti (Fig. 5). Leaf cushions were the dominant tissue con- sumed, accomplished usually by a partial evacuation of in- ner parenchyma surrounding the vascular trace, resulting in expansive galleries interconnected by occasional tun- nels (Pig. 5). Other lycopsid tissues that were consumed include the distal laminae of cones, roots and outer stem periderm. Two examples of borings in calamitean wood and one in a Botryopteris fern were noted. Parenchyma in Lepidophloios leaf cushions was the fa- vored tissue of wood-boring mites during the Early Penn- sylvanian and early Middle Pennsylvanian in the Appala- chian Biisin (Figs. 3; 6a,b,d-h). This host preference also has been documented from the later Middle Pennsylva- nian Mineral Coal-BaU Flora of Kansas (Baxendale, 1979). In both the New Castle and Upper Path Fork coal- ball floras, lycopsids are the dominant taxic constituent, although cordaites approach codominance in the Upper Path Fork {Table 1). Although representing 36.7 percent of the Upper Path Fork flora, only 7.1 percent of the docu- mented feeding traces involved cordaites. Thus, despite rather similar contributions to biomass, lycopsid tissues were attacked about nine times more frequently than those of cordaites (Fig. 6). Upper Path Fork lycopsids, rep- resenting 54.5 percent of the flora, accoionted for 62.9 per- cent of borer feeding traces. The underutilization of cor- daites (though see Fig. 6i) is contrasted to greater frequen- cy of detritivory in sphenopsids, ferns, and to a lesser de- gree seed ferns, each of which constitutes between 2 and 4 percent of the flora but between 4.3 and 12,9 percent of specimens with feeding traces made by mite borers. How- ever, it is possible that this absence oiF cordaitean detriti- vory may be more apparent rather than real, since in oth- er coal ball floras cordaitean wood is comminuted into un- recognizable detritus. One of the earhest examples of cop- rophagy in Euramerican coal-swamps, demonstrating the secondary consmnption of tissues, is from the Upper Path Fork Coal-Ball Flora {Fig. 6c). The Rock Spring Coal-Ball Flora, also from the central Appalachian Basin, contains a similar 1.5 ratio of domi- nant lycopsids {50.9 percent) to subdominant cordaites (33.0 percent). Both the Upper Path Fork and Hock Spring coal-ball floras possess low shoot-to-root ratios (Table 1), indicating a greater contribution fi?jm below-ground tis- sues than from aerially-deployed tissues, includingtrunks (Phillips et al., 1985). This absence of canopy-derived tis- sue suggests substantial degradational loss of litter, and is contrasted to the preponderance of below-ground structur- al tissues illustrated in Figs. 7 and 8, wherein tunneling detritivores attacked tissues ranging from apparently sol- id, undecomposed wood to punky tissues that were soft. (See Fig. 7a for end-members of this continuum.) Unlike the Upper Path Fork Coal-Ball Flora, Rock Spring corda- ites are the principal taxic constituent consumed by detri- tivorous borers (but see Fig. 8e), comprising 33.0 percent of the plant biovolume but 55.9 percent of the attacks by borers, indicating for the first time that the approximate proportional consumption of lycopsid tissues differed. Al- 330 LABANDEIRA ET AL TABLE 2?Diameters of paleozoic coproiites attributed to oribatid mites. A. Pennsylvanian Ag? ; coal-swamp floras from North America (this study). UIUC Plant tissue context Statistical characterization' Maxi- Mini- Coal-ball flora UIUC peel slide of coproiites mum mum Mean Variance N New Castle 38374-Btop 22,400 Lepidophloios leaf cushion parenchyma 61 135 95.5 18.117 58 Upper Path Fork 28729-Btop 22,726 Lepidophloios leaf cushion parenchyma 62 104 83.1 8.381 64 Rock Spring 32479-A 22,446 Cordaitean wood 43 58 49,8 3.782 42 Rock Spring 32479-Ctop 22,403 Cordaitean wood 41 64 50.2 5.390 54 Herrin 3807?-Etop 22,612 Myeloxylon (medullosan) ground tissue 63 39 53.0 7.884 32 Herrin 38030-Ktop 22,515 tissue fragments in macrocoprolite 60 38 47.3 5.670 41 Herrin 38066-Ctop 22,354 tissue fragments in macrocoprolite 40 63 53.7 5.771 27 Herrin 38035-Hbot 22,512 Akthopteris palisade parenchyma 35 61 49.2 6,484 39 Baker 37326-Gbot 22,492 Sphenophyllum. secondary phloem 44 81 59.6 7.866 49 Calhoun 10765-Cbot 22,670 Psaronitis inner root mantle cortex 70 148 106.6 17.614 80 Calhoun 31024-Btop 22.329 Psaronius root parenchyma 55 100 71.2 12.925 46 B. Previous Studies. Plant tissue Diameter Flora and age Locality context of coproiites (p-rn) Reference Esnost (Middle Mississippian) Esnost and Roanne, Botryopteris, fern stem 70^100 (Rf ix & Galtier, 1986; Rex, France ] L986) Upper Path Fork (Kashirki- Cranks Creek, Ken- "Premnoicylon", cordaitean wood c. 40-55= (Cichan & Taylor, 1982) an) tucky Hamlin (Kashirkian) Lewis Creek, Ken- "?Premnoxyhn", cordaitean wood, 30-60, 50 (Taylor & Scott, 1983; Scott tucky other tissues & Taylor, 1983) Mineral (Podobkian) West Mineral, & Lovi- Cordaites leaf mesophyll; Lepido- 3?M0 (Baxendale, 1979) lia, Kansas phloios leaf cushion parenchyma Rotliegende (Lower Permian) Wetterau, Germany Dadoxyhn, probable cordaitean 20-30 (Gioth & Wilde, 1992) wood ' Maximum, minimum and mean values given in micrometers (fxm). =* Estimated from ovoidal and ellipsoidal shapes of coproiites in photographs and stated lengths of approximately 75 pm. though lycopsids constitute about half of the biomass, they only represent 21.8 percent of borer feeding traces. Asso- ciated with these data is the observation that, since pre- served cordaitean and calamitean tissues consist domi- nantly of secondary xylem (Costanza, 1983), the geometry of the borings changed to long, mostly unbranched tunnels that parallel the wood grain and only occasionally expand into galleries (Figs. 4f,g; 7; 8a-d,g). This style of tunnelir^ contrasts with the expansive galleries in leaf cushions that are occasionally connected by narrow tunnels in the New Castle and Upper Path Fork coal-ball ?oras (Figs, 3; 5; 6a,b,d-h). The extensiveness of tunnelii^ in Rock Spring cordaites reveals three types of borii^s: (i) tunnels with dense accumulations of coproiites that lack buildup of comminuted but undigested wood fragments (Figs. 7b- f; 8g), (?) tunnels with well-formed coprohtes amid signif- icant accumulations of unprocessed and indistinctly lay- ered wood fragments (Fig. 8a,b), and (iii) rare tunnels con- taining distinctly stratiform accumulations of conuninut- ed frass with poorly-defined coproiites (Fig, 8c,d), Since xylophagotis mites exhibit a variety of feeding styles in modem woods, these probably represent traces of differ- ent oribatid mite taxa. The Murphysboro Coal-Ball Flora is of Middle Pennsyl- vanian age, and consists dominantly (75.0 percent) of ly- copsid biomass (PhiUips et al., 1985). Compared to previ- ous coal-bail floras, the proportional contribution from cor- daites was more than halved, to 14.5 percent. Other major taxa represent less than 11.? percent of the flora. Al- though lycopod abundance and frequency of consumption by mites are approximately 4.4 times greater than that of cordaites, both taxa seem equal in detritivore preference, since their proportional contributions to total incidence of detritivoiy is similar to thefr respective abundances (Ta- ble 1). The younger Herrin Coal-Ball Flora, from the Illi- nois Basin, similarly contains a high biomass value for ly- copsids (68,9 percent), although the incidence of boring into lycopsid tissues (F^s. 9; lie) tracks lycopsid domi- nance at a lower level of 52.8 percent. The proportional representation of ferns (Fig. lOh), seed ferns (F^. lOa-g), and sphenopsids increases significantly, however, and is roughly congruent with the incidence of tissue boring. Cordaites represent only 0.5 percent of the total plant bio- mass but account for 16.6 percent of the incidence of detri- tivore consumption. Notably, there is evidence for borers attacking a broad spectrum of organs and tissues in host plant taxa, including for lycopsids not only leaf cushions (Figs. 9a-c; lie), stem wood, and lateral laminae of cones (Figs, 9d-e), but also megasporangial wall (Figs. 9f,g) and cone axes. There is mii3?nal evidence for mite consump- tion of pendermal tissues. Locally abundant fohar mate- rial also provides evidence for penetration by borers (Figs. 10 a-d); stem and petiolar tissue of medullosans and ferns demonstrate the selective penetration of sclerenchyma and parenchyma by small borers (Figs. lOf^i). Like lycop- sid megaspores, large meduUosan seeds with thick and in- durated integuments occasionally exhibit invasion by ar- thropod detritivores (Fig. lOe). It is also significant that ORIBATID MITES IN COAL-SWAMP FORESTS 331 ,.^' ? -.-.?iSSSSS?^m FIGURE S?Mite consumption of leaf-cushion parenchyma in the lycopsid Lepidophloios harcouriii, from the New Castle Coal. (Winston and Phillips, 1991). All photographs are from slide 22,400 (peel specimen 38374-Btop). Solid scale bars =1.0 mm; barred scale bars = 0.1 mm. (a) Oblique tangential section of leal cushions from a segment of a LepidophioiosVMig, some with galleries excavated by mites, (b) Enlargement of distal region of leaf cushion indicated at up per-left of (a), showing gallery and constituent cc prolites in parenchyma, (c) Enlarg m?nent of gallery outlined in (b). (d) Enlargement of basal region of leaf cushion indicated at central-right in (a). microdetritivore coprophagy has been documented in greater amounts (Figs, lla-d) than in any previous coal- ball flora. This general pattern of major consumption of di- verse lycopsid tissues and a substantial representation of detritivory in the other plant groups occurred in the youn- ger Baker Coal-Ball Flora (Table 1; Figs, llf.g). After the demise of lycopsid-dominated forests at the end of the Middle Pennsylvanian, Psaronius tree-ferns be- came the dominant vegetation in the equatorial Euramer- ican wetlands until the Early Permian. In fact, Psaronius and other ferns account for 64.3 percent of the plant bio- mass, and 76.7 percent of feeding traces by borers. Lycop- sids, dominant in both of these respects during the Early and Middle Pennsylvanian, shrink to 1.9 and 5.5 percent, respectively. Seed ferns are ranked second (24.2 percent) in overall biomass, and contribute 15.1 percent of the feed- ing traces hy detritivores. Thus the character of early Late Fernisylvanian coal swamps shifted to forests dominated by ferns and seed ferns, collectively accoimting for approx- imately 90 percent of both biomass and consumption by detritivores. The peats resultir^ from these forests were penetrated extensively by roots, many of which contain timnels with coprohtes (Figs. 12a,c-e). Generally, in con- trast to preserved Middle Pennsylvaman coal-ball floras, there was considerably more detritivore consumption of fohar and reproductive tissues (Figs. 12cXg)> probably at- tributable to a relative paucity of structural tissues, es- pecially wood. The aerial and subterranean root manties o?Psaroniiis trunks were a limited but conspicuous excep- tion, showing evidence of tuimeling within individual root elements (Pig. 12b). These observations undoubtedly re- flect the decline of major woody cordaites during the late Middle to Late Pennsylvanian (Phillips et al., 1974; Phil- Ups and Peppers, 1984; DiMichele and Hook, 1992). More- 33S LABANDEIRA ET AL FIGURE 6?Tissue types, including foliar parenchyma, secondary xylem, and various macrocoprolitio fragments, of lycopsids and cordaites consumed by mites in the Upper Path Fork Coal. Soiid scale bars = 1.0 mm; barred scale bars = 0.1 mm. (a,b,d to h) ; Radial and tangential sections of of leaf cushions from the lycopsid Lep??ophioios h?rcourtii, shown with mite coprolites. (a) Radial section of an axis bearing five leaf cushions that exhibit excavation of leaf-cushion parenchyma. Slide No. 22,727 (peel 28686-A). (b) Enlargement of gallery region in one of the leaf cushions outlined in (a), (c) Tunnel, indicated by white arrow, containing mite microcoproutes within a macrocoprolite, evidence for coprophagy. Slide No. 22,438 (peel 28933-?iop). (d) Tangential section of axis bearing nine leaf cushions with coprolite-containing galleries. Slide No, 22,433 (peel 28892-Btop). (e) Enlargement of leaf cushion at lower-left in (d), exhibiting three major galleries, (f) Enlargment of leaf cushion at upper-left in (d), exhibiting two galleries connected by a tunnel, (g) Radial section of Lepidophloios haivoutii\eat cushions containing galleries of excavated parenchyma. Note empty regions in the distal part of several leaf cushions and replacement by clusters of small, isodiametric coprolites. (h) Enlargment of distal leaf cushion region in (g), showing the distribution of cop rollte clusters at the margin of the central gallery. Slide No. 22,726 {peel 28729-Btop), (i) Tunnel containing microcopfelites in secondary xylem of the cordaitean root, Amyelan. Slide No. 22,428 (peel 28799-?top). ORIBATID MITES IN COAL-SWAMP FORESTS 333 ^_^,^:i'-^. '%-"^ ' .-a ??*? ??- FIGURE 7?Mite consumption of wood from the Rock Spring Coal. Solid scale bars = 1.0 mm; barred scale bars = 0.1 mm. (a) Transverse section of two pieces of cordaitean wood, shtowing juxtaposition of undegraded cordaitean secondary xy?em below, and mite-riddled, punky, wood above. Slide No. 22,446 (peel 32479-A), (b) Transverse section of cordaitean wiood, sfiowing circular to elliptical cross-sections of mite tunnels in secondary xylem. UlUC slide 22,403, from UIUC peel 32479-Ctop, (c) Cross-sections of mite tunnels from a transverse section of cordaitean wood. Slide No. 22,441 (peel UIUC 32479-Bt30t>, (d) Transverse to oblique section o? three mite tunnels in cordaitean wood, showing variation in coprolite size that indicate instar stages. Slide 22,403 (peel 32479-Ctop). (e) Enlargement of circular tunnel showing the composition, shape, and texture o? mite coprolites, probably in cordaitean wood. Slide No. 22,406 (pee! 32330-Bbot). (f) Tangential section of probable calamitean secondary xylem, showing parallelism of wood grain and tunnel orientation. Slide No. 22,406 (peel 32330-Bbot). 334 LABANDEIRA ET AL. --^-i h;:-.:?w>r^--7fc ^ ?^qil S?T'?^d ?l^^;Crs;avi;--#:^ M? If [ " \ FIGURE 8?Mite consumption of tissues in cord alte s, ferns, and macrocoprolltes in the Rocli Spring Coal. Solid scale bars = 1.0 mm; barred scale bars = 0.1 mm. (a) Transverse section of calamitean wood, containing a longitudinal section of a mite tunnel. Frass consists ol ellipsotdat fecal pellets and comminuted wood within the tunnel. Slide No. 22,440 (peel 32350-Cbot). (b) Enlargement of region in outlined in (a), (c) Longitudinal section of a calamitean wood showing a longitudinal section of a mite tunnel with a characteristic layered pattern of frass and lack of we II-formed coprolites. Slide No. 22,471 (peel 32277-Dtop). (d) En la rg ment of of region delineated in (c), showing the layered frass. (e) Cross-section of a foliar memt?er of the filicalean fern, Botryoptehs, displaying in outer ground tissue a tunnel containing a cluster of coprolites. Slide 22,673 (peel 32274-A3-side 2). (f) A cluster of coprolites in a tunnel occurring within a macrocoprolite. Note the heterogenous tissue fragments with thickened cell walls in the macrocoprolite. Slide No. 22.456 (peel 32277-Dtop). (g) Longitudinal section of a degraded fragment of cordaitean wood, containing a cross-section a mite tunnel and a penetrating root at lower right. Slide 22,672 (peel 32274-15A- 4top). ORIBATID MITES IN COAL-SWAMP FORESTS 335 FIGURE 9?Mite consumption of various lycopsid taxa and tissue types in the Herrin Coa!, from the Illinois Basin. Solid scaie bars = 1.0 mm; barred scale bars = 0.1 mm. (a) Transverse section o? Diaphoroden?ron stem, showing a leaf cushion with evacuated parenchyma and replacement by mite coprolites. Slide No. 22,537 (peel 38012-Dbot). (b) Part of a Lepidocarpon axis, with degraded tissue in the sporophyll. Parenchyma region in the sporophyll now occupied by mite coprolites. Slide No. 22,612 (peel 38070-Etop). (c) Mite excavation of parenchyma in Lepidophloios hallii\eaf cushion. Slide No. 22,517 (peel 38040-Gbot). (d) Longitudinal section of the lycopsid cone tip, ?.ep/docafpoo, showing sporophyll lateral laminae. Note that the basal indurated region of lateral laminae possess galleries filled with mite coprolites. Slide No. 22,524 (peet 38022-ltop). (e) Enlargement of basal regions of lateral laminae outlined in (d). (f) Longitudinal section of a Lep/docarTranmegasporangial wall and lateral laminae. Note mite tunnels and chamber in the basal attachment region to the cone axis. Slide 22,363 {pee! 37112-!top). (g) Enlargement ot region indicated in (i). 336 LABANDEIRA ET AL FIGURE 10?Mite consumption of tissues from seed ferns in the Herrin Coal. All specimens are from Sttawneetown, except (h), from Carrier Mills. Solid sc^e bars = 1,0 mm; barred scale bars = 0.1 mm. (a) Transverse section of Alethopteris pinnule. Slide No. 22,520 (peel 38035- Hbot). (b) Enlargement of central pinnule region in (a), showing excavation of palisade and hypodermis layers, and vascular bundles, (c) Oblique longitudinal section of a Myeloxytan axis, exinibiting vascular strands, intervascular parenchyma, and resin canals. Note outlined vascular strand partly consumed by mites. Slide No. 22,577 {peel 38043-Lbot). (d) Enlargement of vascular strand in (c) with a coprolite- containing mite tunnel, (e) Sclerolesta of ttie medullosan seed, Pachytesta (at bottom), containing a row of mite coprolites (at top) witfiin the nucellus tissue (arrow). Slide No. 22,416 (peel 37112-Etop). (f) The medullosan frond axis, Myeloxylon, showing mite galleries among ground parenchyma and fibrovascular bundles of the frond periphery. Slide No. 22,520 (peel 38035-Hbot). (g) Calamitean wood with a mite tunnel. Slide 22.612 (peel 38070-Etop). (h) Mite tunnel in outer sclerenchyma of a Slipitopteris tree-fem petiole. Slide No, 22,608 (peel 38078-Ctop). ORIBATID MITES IN COAL-SWAMP FORESTS 337 FIGURE 11?Mite consumption of plant tissues from tfie Herrin (a-e) and Balder (f^g) Coais. Ail specimens are from Sfiawneetown, except a and b, which are from Carrier Mills. Solid scale bars = 1.0 mm. (a) Cross-section of a macrocoprolite consisting of digested tissue fragments and two galleries containing mite microooprolites. UIUC slide 22,354, from UIUC peel 38066-Ctop. (b) Enlargement of galleries indicated in (a), (c) Longitudinal-section of nnacrocoprolite with pronounced exposure rind, composed of tissue fragments and microcoprolite-bearing tunnels. Slide No. 22,515 (UIUC peel 38030-Ktop). (d) Eniargement of tunneled region indicated in (c). (e) Obliquely transverse section of leaf cushions from the lycopstd Lepit?ophloios, showing an excavated gallery in a leaf cushion at center, and replacement by cop routes along inner region of epidermal tissue. Slide No. 22,730 (UIUC peel 38017-Ktop). (f) and (g): An obliquely transverse section of a stem of Sphenophyllum, indicating consumption of tissues between wood and periderm. Slide No. 22,492 (peel 37326-Gbot). (f) Entire stem, showing major tissues, (g) Enlargement of region at central-left, delineated in (f), exhibiting secondary xylem to left, microooprolites in center, and phellem to right. 338 LABANDEIRA ET AL. "X' ??j-??v, V'. ?'XV^^-V^L'TI ^^?3;jt4?:^^^ ?. ??^..dl^ ^?SlSPi FIGURE 12?Mite consumption of various plant tissues in fem, sphenopsid, and lycopsid taxa from the Cathoun Coal, Solid scale bars = 1.0 mm; barred scale bars =0.1 mm. (a) Oblique, longitudinal section of a Psaronius root, witti the stele in the center surrounded by microcoprolites in the cortex. Slide No. 22,393 {peel UIUC 30679-Bbot). (b) Occurrence of mite coprolites in the inner root mantle lumina of Psaronius. Slide No. 22,670 {peel 10765-Cbot>. (c) At top, a ^henopsid Annutaria leaf in which the spongy mesophyll has been evacuated and replaced by a cluster of mite coprolites; note avoidance of palisade mesophyll with thickened cell walls. Fecal pellets occur in Psaronius root at bottom. Slide No. 22,667 (peel; 31272-Bbot), (d) Longitudinal section of a Psaronius root exhibiting a centra! tunnel of excavated tissue and replacement by mite coprolites. Slide No. 22,662 {pee! 31272-Ctop). (e) Same as (d), but oblique transverse section. Slide No. 22,329 (peel 31024-Btop). {f) Oblique transverse section of Aletho?tens foliage, exhibiting removal of mesophyll tissue between outer epidermis and inner vascular bundle. Slide No. 22,662 (peel 31272-Ctop). (g) Same as (f), but transverse section of midrib. Slide No. 22,643 (peel 38600-Bbot). OHIBATID MITES IN COAL-SWAMP FORESTS 339 over, the ?cologie radiation of new herbivore functional- feeding groups (Labandeira and Phillips, 1996a, 1996b) may have qualitatively altered those detritivore feeding strategies that survived into the Late Pennsylvanian. DISCUSSION Recently, several studies have evaluated plant damage and coprolite contents preserved va peat litter from the earhest known forested ecosystems (summarized in La- bandeira and Beall, 1990; DiMichele and Hook, 1992; and Scott et al., 1992). These, and our ongoing studies, have demonstrated that arthropods responsible for many feed- ing traces can not be identified with taxonomic precision. Only stereotyped or unique interactions can be assigned to the level of an insect taxonomic order or a more encom- passing, specific clade of insects (Labandeira and Phillips, 1996a, 1996b). In our assessment of the tissue damage documented in this report, we recognize characters of Ufe history and behavior that allow a more precise identifica- tion of the responsible arthropods than previously possi- ble. This evidence supports oribatid mites, particularly the superfamilies Euphthiracaroidea, Phthiracaroidea, Her- mannielloidea, Carabodoidea and Liacaroidea, as the ar- thropod clades responsible for the extensive borings into a diversity of Pennsylvanian plant hosts and plant tissues. Autecological data such as borings, galls, foliage chew marks, and piercing-and-sucking tracks have now been doounented from many Pennsylvanian plants {Labandei- ra and Beall, 1990; Scott et al., 1992; Labandeira and Ph?- lips, 1996a, 1996b), indicating that previous statements, such as "the analysis of Scott and Taylor (1983) carries coprolite evidence as far as it can go" (Shear and Kukalo- v?-Peck, 1990: 1829), were premature. The identities of plant-interacting arthropods probably w?l never be known at low taxic levels because most of the Paleozoic ar- thropod fauna was replaced by modem lineages after the Permo-Triassic extinction (Labandeira and Sepkoski, 1993; Seiden, 1993a; Labandeira, 1994). Although Paleo- zoic trace fossils may suffer from poor taxonomic resolu- tion, the taxic and ?cologie data they represent are ontologicaUy separate. The latter are in fact high-resolu- tion data on the geochronological timing of fUnctional- feeding-groups and dietary guilds in Paleozoic ecosystems. Oribatid Mites; Fabricators of the Borings Organisms producing fecal peUets and occurring in ste- reotyped tunnels within decomposing plants can be deter- mined with greater specificity and confidence than can those organisms excreting larger fecal peUets that occur in isolation within ambient soil litter, without an obvious plant-host association. Because of this greater potential for identification, we have specified five discrete lines of evidence for taxonomic assignment of these coprohtes. They are (1) coprohte size, (2) coprolite shape, (3) coprohte surface texture, (4) coprohte contents, and (5) surrounding plant tissue context of the coprohte. When these five cri- teria are applied to the extensive Literature on fecal pellets and tunneling behaviors produced by modem terrestrial anneUds and arthropods, several groups are obvious can- didates as producers of the coprolites and tuimels de- scribed in this report. They are oligochaete annehds, in- sectan microarthropods, immatiu-es of myriapodan and insectan macroarthropods, collembolans, and oribatid mites. Our evidence strongly indicates that oribatid mites were the producers of these distinctive coprohtes within plant tissues. Enchytraeid, Iximbricid and other oUgochaete annehds are common and numerically abundant constituents in or- ganic rich so?s and decomposing litter (WaUwork, 1976a, 1976b). Although ohgochaetes consimae oi^anic soils rich in plant material, they rarely ingest solely plant detritus. Their fecal peUets occur in the lower F and especially H horizons, consist of a dark and nearly homogenous mix- ture with scattered minerafic matter, and have surfaces with considerable reUef (Kubiena, 1955; Zachariae, 1965; Rusek, 1975; Ponge, 1988). The peUets also are rarely en- countered intact, and have a tendency to disaggregate (Jongerius, 1963; Babel, 1968). Oligochaetes have fossil records extendii^ to the Oligoc?ne (Wills, 1993), but are not known to tunnel or otherwise subsist on wood diets (O'Connor, 1967). Similarly, fecal peUets from various ex- temaUy-feeding insectan microarthropods, such as diplur- ans, archaeognathans and thysanurans, are not found within plant-host tissues, nor are they composed princi- paUy of ingested wood fragmente. AdditionaUy, archaeog- nathans and probably thysanurans have irregularly shaped fecal pellets with ragged edges characterized by projecting firagments (Hartnack, 1943; D.H. Headrick, pers. comm.). Some smaU adult insects are wood borers (Frost, 1959; Hickin, 1975; Eaton and Hale, 1993), but do not produce fecal peUets in the minuscule size ranges of the coprohtes described herein. The smallest adult xyloph- agous insects?termites, bostrychid and scolytid beetles, and carpenter ants?produce fecal peUete of different shapes and significantly larger sizes than those described from Pennsylvanian plant tissues (Eckstein, 1939; Lin- sley, 1943; Weiss and Boyd, 1950; Simeone, 1965; Eaton and Hale, 1993). Subadult instars of myriapods and insecte produce fecal peUets that occur in litter and in association with five plant tissues. Some subadult instars of diplopods produce fecal pellete approaching the si2e range of oribatid mites, although their shape difiers significantly by being elon- gate (often two or more times the diameter), possessing an irregular and bumpy surface, and containing well-defined and often identifiable fragmente of plant tissues (Zachar- iae, 1965; Paulusse and Jeanson, 1977). Although myria- pods were present dirrir^ the Late Paleozoic (Almond, 1985), and although certain species are frequently found in decaying wood (Wallwork, 1976), miUipedes are unlike- ly to have caused any of the traces seen in our study. When feeding as endophages in woody materials, they create rel- atively large chambers, rather than narrow tunnels. By contrast, many insect subadulte?particularly endoptery- gote larvae of the Cole?ptera, D?ptera, Lepidoptera, and Hymenoptera?are wood borers and produce copioiis quantities of fecal pellets containing fragmente of wood and other indurated plant tissues (Hickin, 1975; Mamaev, 1977; Crowson, 1981; Eaton and Hale, 1993). However, fe- cal peUet shapes of endopterygote larvae are fimdamen- taUy different than aU other wood-boring arthropods: they consist of cylindrical segmente of an extruded bolus mass whose ends indicate brittie fracture or other types of 340 LABANDEIRA ET AL. TABLE 3?Published descriptions of modem oribattd mite and Ct?lemboian hexapod fecal pellets. Pellet size Taxon Stage (diameter x length) Pellet shape Data source Oribatid mites Ameronothridae m 45X eSuJtnto 100 X 150 M-m 50 X 60 tim elliptical, spheroidal Schulte, 1976 Nothridae Nothrus bicilitztus (Kach) larva protonymph deutonymph tritonymph adwlt 27.3 X 40.6 fim 37.1 X 61.6 tim 52.5 X 87.5 tJLm 70.7 X 113.4 um 93.8 X 176.4 jjim not mentioned Saichuae, et al-, 1972 Phthiracaiidae Phthirnmrus ferrugineus nymph 57 X 85 M-m spheroidal and Riha, 1951' (Koch) adult 100 X 160 liJ? ovoidal Phthiracaridae Rhysotritia duplicata unknown 140 X 220 (j,m spheroidal and Bal, 1968 Grandjean Microtritin mijiima unknown 7 X 12 (jon ovoidal Bal, 1968= (Berlese) Mropacarus striculus unknown 34 X 57 )im to Ral, 1968^ (Koch) 75 X 100 Jim Oribatida m 40 X 100 \i.m in length ?ovoidal Drift, 1964 Oribat?da m 50 X 90 \im to 120 X 260 \i.m not mentioned Bal, 1970 Oiiba?da m 30 to 150 urn not mentioned Babel, 1975 Oribatida m 30 to 50 Jim in diameter ovoidal, spheroidal Rusek, 1975 Oribat?da n 140 X 200 ii.m ovoidal, spheroidal Eusek, 1984 Collembolan hexapods CoUembola ?a less than 50 (xm not mentioned Jongenus, 1963 CoUembola m 50 to 200 (1,111 in diameter ?ovoidal Drift, 1964 Collembola m 30 to 90 (>100) p.jn spheroidal Kuaek, 1975,1985 CoUembola (probably) ?u 30 X 50 tun to 90 X 125 fxjn not mentioned Rusek, 1975 CTollembola aU 50 to 180 (un not mentioned Babel, 1975; Pawiuk, 19S5 ' As Phtkiracarus ligneus W?lmann. * As Rhysotritia minima (B?rlese). ^ As Steganacarus striculus (Koch). break^e. These pellets often have six longitudinal fur- rows that are a consequence of six longitudinal muscles that contour the passing food bolus in the rectum (Grim- stone et al., 1968). Thus the cylindrical fecal pellet shapes of endopterygote larvae are a consequence of continuous or periodic extrusion, and differ from the spheroidal to e?- hptical shapes in other terrestrial wood borers that result nxim individual and periodic packaging (Weiss and Boyd, 1950). For these reasons, ohgochaete annehds, insectan microarthropods, and the subadult instars of myriapods and insects are excluded as candidates responsible for pro- ducing the endophytic coprolites documented in this re- port. With regard to collembolans and oribatid mites, there is documentation of microarthropod fecal pellet size and shape in ground litter and peat-rich soils of modem north- temperate forests. Available records, compiled in Table 3, indicate that ovoidal fecal pellets, approximately from 3? X 40 Jim to 140 X 200 jim, and spheroidal pellets from 30 to 200 fjLm in diameter, are common to very abundant in litter and upper soil horizons, and invariably originate from oribatid mites and collembolans. While their fecal pellets almost completely overlap in size, their shape and surface structure difiter, Oribatid mite fecal pellets bear relatively smooth surfaces {Tarman, 1968; Cervek, 1976) whereas collembolan fecal pellets are variously textured and irregular, with tissue fragments projecting beyond the general surface relief (Zachariae, 1963, 1965; Rusek, 1975). These distinctions of fecal pellet surface rehef are related to the peritrophic membrane, present in mites (Woodring and Cook, 1962; Dinsdale, 1975; Evans, 1992), but apparently absent in collembolans, resulting in smoother surfaces of mite pellets. Additionally, differing mechanisms of processing food and perhaps diet in the two microarthropod clades contribute to differences in surface texture. Although there are some taxa in both groups that ORIBATID MITES IN COAL-SWAMP FORESTS 341 exhibit dietary convei^ence, particularly on microfiir^, most detritivorous oribatid mites prefer plant tissues with high cellulose content, including sclerenchyma and wood (Pande and Berthet, 1973). Collembolans consume softer tissues, preferring more digestible substrates such as bac- teria, algae, fungi, partly degraded parenchyma, pollen and spores, fecal pellets, and dead animals (MacNamara, 1924; Poole, 1959; Chiistianson, 1964; Adams and Salm- on, 1972; Schulte, 1976; Haq, 1982; Aitchison, 1983; Tak- eda and Ichimura, 1983). Selectivity is poorly developed in most collembolans (Christianson, 1964; Singh, 1969; Veg- ter, 1983; but see Shaw, 1988), and there is no indication that any species regularly subsists on wood or a similar ceUulosic or ?gnified tissue. As major microarthropod decomposers that have been documented in north-temperate ecosystems, oribatid mites physically reduce Htter into particvdate debris by consumption of a diverse spectrum of plant tissues. This detritivory is accomphshed by unique mouthparts and chelicerae that are modified for finely comminuting plant material (Schuster, 1956; Wallwork, 1958; Dinsdale, 1974; Schulte, 1976; Haq, 1982), Adults and subadults in some groups bore through hard plant tissues such as wood, bark, and fibrovascular bundles, whereas those of other groups choose softer parenchymatous tissues in conifer needles (Fig. 4c-e), angiosperm broadleaves (Fig, 4a), roots, and other plant organs (Woodring and Cook, 1962). Thus tissue-boring oribatid mites are subdivided into three dietary types: (i) varied surface litter, mostly leaves that contain spongy parenchyma; (ii) wood (xylophagy), indudir^ taxa that commonly bore into leaf petioles, cone scales, or other indurated tissues; and (iii) pre-existing di- gested plant material found as arthropod fecal pellets (coprophagy), Collembolans, by contrast, lack appropriate mouthpart structure for boring into plant tissue (Goto, 1972). Although a few coUembolan species bear mandibu- lar incisors that can be used to tease apart woody tissues (MacNamara, 1924; Wolter, 1963), none are known to cre- ate tunnels or galleries by their feeding actions. The process and consequences of oribatid mite con- sumption of plant tissues has been elucidated by studies of the immediate biologic fate of newly-fallen htter and the int^ration of degraded litter into tie upper layers of or- ganic sous (Butcher et al., 1971; Webb 1977; Swift, 1977; Seastedt, 1984), Historically, the best documented exam- ples of litter degradation are (i) tunneling into conifer nee- dles and, to a lesser degree, petioles of angiospermous leaves, (ii) surface skeietonization of angiosperm leaf blades, (iii) tunneling into conifer and dicot wood, and (iv) feeding on macroarthropod fecal pellets containing plant tissues. Two methodological approaches have been used to address the degradation of above-ground plant tissues? autecologieal studies emphasizing the role that species or individuals have in consuming particular plant tissues, and synecological examinations that evaluate multitroph- ic feeding strategies of microarthropod communities in ht- ter breakdown. Oribatid mites gain entrance into conifer needles by producing surface depressions (Harding and Stuttard, 1974) that range from 400 p-m in diameter (documented by Hartenstein, 1962c) to holes that are substantially smaller (Figs. 4c-e). These mites subsequently endophagously consume mesophyll parenchyma and softer tissues, leav- ing vascular bundles and epidermal cuticle intact (Jacot, 1939; Dinsdale, 1974; Babel, 1975; Gourbiere et al., 1985, 1987; Lions and Gourbiere, 1988,1989), Once conifer nee- dles have been processed lay oribatid mites in the L and upper F layers (Pig. 4b), they essentially consist of elon- gate sacs containing fecal pellets and occasional fragments of nondigestable tissue (Kubiena, 1955; Jongerius, 1963; Zachariae, 1965; Hal, 1973; Rusek, 1975). These sacs are eventually ruptured by macroarthropod and other inver- tebrate bioturbators, and become incorporated into deep- er, organic-rich soil horizons. By contrast, planar angio- sperm leaves are frequently skeletonized by detritivorous insects and particularly ectophagous and endophagous or- ibatid mites, resulting in intact primary and secondary veins and often a tertiary vein meshwork in more impal- atable species (Zachariae, 1965; Bal, 1968,1982; Harding and Stuttard, 1974; Rusek, 1975; K?hnelt, 1976). Tissue consumption of Pennsylvanian fohar parenchy- ma ranges from lycopsid leaf cushions (Figs. 5; 6a-e; 9a,b), to seed fern pinnules (Figs, lOa-d, 12f,g). These tissues are structurally identical to those documented in mite- consumed parenchyma of modem plants (e.g., Kubiena, 1955; Jongerius, 1963; Kubikova and Rusek, 1976; Babel, 1975; Cohen and Spademan, 1977; Cohen et al., 1989). Presently, there is a broad spectrum of oribatid mite taxa associated generally with consumption of parenchyma- tous tissues, including constituent subtaxa that commonly ingest leaf utter but are typicaUy associated with wood, such as phthiracaroid, euphthiracaroid, and cepheoid mites (Pande and Berthet, 1973; Niedbala, 1992), Based on differences in foss? tunnel contents and geometry men- tioned previously, it is highly likely that a diversity of ori- batid mite taxa also were associated with the consumption of diverse fohar material during the Pennsylvanian. The wood-boring life habit has originated several times within oribatid mites, occurring not only in the omnipres- ent box mites of the Phthiracaroidea and Euphthiracaro- idea (Michael, 1882; Niedbaia, 1992), but also in some members of the Epilohmanniidae (Norton, unpublished), Lohmanmidae (Haq, 1984; Haq and Konikkara, 1988), HermanieUidae (Michael, 1882; Riha, 1951; Schuster, 1956; Wallwork, 1967), Carabodidae (Michael, 1882; Riha, 1951; Wallwork, 1967), Xenilhdae (Michael, 1882; Norton, unpublished), Liacaridae {C?ourbi?re et al., 1985,1987; Li- ons and Gourbiere, 1988,1989) and Oribatuhdae (Woodr- ing and Cook, 1962; Wallwork, 1967). Virtually all of the textual and photographic documentation of wood boring in modem oribatid mites historically is of phthiracaroids and euphthfracaroids, particularly the prominent genera Phthiracarus and Steganacarus of the Phthiracaridae and Oribotritia of the Oribotrit?dae (Jacot, 1936,1939; Riha, 1951; Schuster, 1956; Wallwork, 1957; Haq, 1982; Ponge, 1988; Soma, 1990). Typically phthiracarid mites construct tunnels that parallel the grain of punky to recently fresh wood (Figs, 4a-e), and are circular in cross-section (Fig. 4e), usually of somewhat larger diameter than body width. Originating irom the main tunnel in some species are side branches that are constructed by nymphs and enlarge dis- tally (Wallwork, 1957), Ovoidal fecal pellets are packed within these tunnels, either as occasional clusters occupy- ing the entire tunnel diameter (Riha, 1951; Wallwork, 1957; Drift, 1964), or as a rim of pellets plastered to the tunnel wall (Riha, 1951), or singly on the floor of the tun- 342 LABANDEIRA ET AL nel, As in fecal pellet size, diameters of tunnels vary with taxic affiliation and instar of the mite borrower; some- times an entire ontogenetic sequence is preserved in the length of a tunnel of increasing diameter (Figs. 7d; 9g), reflecting a laiTa->protonymph->deutonymph->trito- nymph->adult sequence (Wallwork, 1957). In modem, north-temperate, forested ecosystems, virtually every iype of wood or similar indurated tissue is attacked by or- ibatid mites, including conifer wood (Drift, 1964; Andr? and Voegtlin, 1981; Ponge, 1988), dicotyledonous angio- sperm wood (Kubiena, 1955; Bal, 1968; Kub?kov? and Eu- sek, 1976), bark (Schuster, 1956; Wallwork, 1957; Harding and Stuttard, 1974), seed testas (Harding and Stuttard, 1974; Harding and Easton, 1984), ovuliferous scales of pine cones (Webb, 1978, 1989), roots (Jacot, 1936; Drift, 1964; Bal, 1968), and even shoreface wrack (Haq, 1984; Haq and Komkkara, 1988). This modem spectrum of tis- sue and organ types parallels the breadth of mite detriti- vory in Pennsylvanian coal-swamp forests. In one of the few studies of a site-specific oribatid mite Community occurring within wood, Wallwork (1957) docu- mented the presence of tritrophic interactions. In decay- ing and moist birch (Betula) branches and twigs m surface litter, four species of xylophages that dwelled in the sec- ondary xylem and lenticels were accompanied by six ad- ditional species coprophagous on the xylophage fecal pel- lets, with both groups occasionally consumed by three co- existing, predatory species. The presence of analogous taxa in Baltic amber (Sellnick, 1919,1931; Appendix 1) in- dicates that the antiquity of similar communities based on wood-consuming mites extends minimally to the Early Ce- nozoic. Increasingly, acarologists and soil biologists have docu- mented the importance of secondary consumption of de- graded plant tissue in the form of coprophagy. Although oribatid mite coprophagy occurs among nymphs subsist- ing on the fecal pellets of older, conspeciiic instars (Wall- work, 1958,1967; Haq, 1984), it is most common in species consuming macroarthropod fecal pellets, including those of millipedes and insects (Schuster, 1956; Nicholson, et al., 1966; Bal, 1973). In fact, coprophagy may be a strategy for detritivorous mites that are more likely to encoimter de- graded, higher plant material and a surface microbial bi- ota on ripened fecal pellets than in surrounding, undigest- ed plant tissues. As a dietary strategy, coprophagy of ma- croarthropod fecal pellets was well-established during the Pennsylvanian (Figs. 6c; 8f; lla-d), indicating that sec- ondary detritivory and consumption of microbes may have been a significant source of nutrition. With the exception of two apparently mycophagous spe- cies of Middle Devonian mites (Norton et al,, 1988,1989), the body-fossil record of oribatid mites is unilluminating prior to the Jurassic (Appendix 1). We know that some or- ibatid mite lineages extend to the Devonian (Norton et al,, 1988), but certainly by Jurassic times the known oribatid mite fauna essentially was modem in taxic aspect (Krivo- lutsky and Druk, 1986). While the body-fossil record of or- ibatid mites is lacking during the Carboniferous to Trias- sic, the trace-fossil record of oribatid mites conversely is present during this interval. This window of well-pre- served, ecological data provides complementary data on the life habits and behavior that contrasts to our lack of knowledge of oribatid mite morphology or systematics. From this window of fossil borings in plants, modem pat- terns of tissue consumption are indistinguishable from that of the Pennsylvanian fossil record of coal-swamp plants. Wood-boring behavior and xylophagous diets al- ready were well developed in woody and other indurated tissues (Figs. 7; 8; 9f,g; lOg), resembling in detail modem oribatid mite borings in gymnospermous and angiosper- mous wood (Wallwork, 1957; Drift, 1964: figure 15; Kub?- kov? and Rusek, 1976, Rusek, 1985). Pennsylvanian-age plant tissues tunneled by oribatid mites include promi- nently cordaites (Figs, 6i; 7; 8a-d,f,g), but also consumed was the trunk-surrounding root mantle of ps?ironiaceous tree ferns (Fig. 12b), leaf cushions, megasporangial wall or lateral laminae of lycopsids (Pigs. 5; 6a,b,d-h; 9f,g), andfi- brovascular bundles within the trunks of mediillosan seed ferns (Pigs. llc,d,f,g). This pulse of trace-fossil data indi- cates that a diverse spectrum of Pennsylvanian-age plant hosts and tissue types were consumed by oribatid mite de- tritivores, suggesting that at least modest taxic diversity was already associated with widespread ?cologie diversity. Contrasts in Wood Degradation from the Late Paleozoic to the Recent The pattern that emerges fi:x)m our study of Pennsylva- nian plant borers and an examination of documented tis- sue-borii^ activity in the fossil record is provided in Fig. 1. The two obvious temporal concentrations of wood-borii^ are the Pennsylvanian for oribatid mites, and the later Mesozoic and Cenozoic for insects. These two dense occur- rences result from differing geologic and taphonomic set- tings of oribatid mite and insect trace fossils, as well as the relative recency and, hence, greater likelihood for strati- graphic completeness of the insect trace-fossU record. The pulse of oribatid mite borings found in Carboniferous coal-swamp deposits is associated with liirge, regional ac- cumulations of coal and permineralized peat; by contrast, later Mesozoic and Cenozoic fossil insect borings occur in plants from varied, mesic habitats, such as fiood-plain for- est and warm-temperate forest of lake margins. Conse- quently, in addition to the puU-of-the-recent mentioned above, the later Mesozoic and Cenozoic records of insect borings represent a considerably broader ecological spec- trum of plant commimities not associated with burial of large volumes of peat in unique habitats, such as those represented in Pennsylvanian deposits. Strengthening the environmental explanation of this pattern is that Pa- leozoic and post-Paleozoic compression floras are pre- served in a similar way (TimeU, 1962; CoUinson et al., 1994). These and other qualitative distinctions in the ori- batid mite and insect records of borers are provided in Ta- ble 4. Dramatic ?cologie, taxic, and geologic contrasts are evi- dent between the trace-fossU records of oribatid mite and insect borers. The ?cologie breadth of oribatid mite detri- tivores has survived admirably to the present, consuming virtually all types of dead plant tissue. For the Early and Middle Pennsylvanian, up to the demise of the lycopsid- dominated coal-swamps at the end of the Middle Pennsyl- vanian (PhUhps et al. 1974, Phillips and Peppers, 1984), einrent evidence supports detritivory as the basis for con- sumption of not only hard plant tissues such as wood, bark, and sclerenchyma, but also most sofl^r tissues. Dur- ORIBATID MITES IN COAL-SWAMP FORESTS 343 TABLE 4?Comparison of the ctocumented trace-lossil records of oribaiid mite and insect borers. Trace-fossil record Feature Oribatid mite borers Insect borers Plant host taxonomic group: Tissues attacked; Location of attacked tis- sues: Fossil occurrence: Regional taphonomic cor- relates Geochronology: Lycopsids, calamites, ferns, cordaitean conifers, pteridosperms Dead tissues ranging from fresh (but fungaUy attacked) to punl^ detritivory At or below ground level Coal swamps Associated with large, regional accumu- lations of reduced plant material Carboniferous Pteridosperms, cycadophytan and coniferan gymnosperms, angiospenns Dead tissues; but more importantly Uve tissues, especially bast {phloem and cambium); detritivory + herhivory At, below, and especially above ground level Subtropical to temperate mesic forests Not associated with large, regional accumulations of re- duced plant material Late Triassic to Recent ii^ the Late Pennsylvanian, this widespread mite detriti- vory was supplemented in the same environments by new types of insect herbivores occurring within the forest can- opy not associated with consumption of hard tissues (La- bandeira and Phillips, 1996a, 1996b). This transition oc- curred when psaroniaceous tree ferns achieved ecological dominance in lowland wetlands across much of the Eura- merican equatorial belt. These new insect herbivores oc- cupied several functional feeding groups during the Late Pennsylvanian, including external foliage feeding, pierc- ing-and-sucldng, gallii^, and sporangivoiy (Labandeira and Beall, 1990; Lesnikowska, 1990; Labandeira and Phil- lips, 1992, 1996a, 1996b; Labandeira et al., 1994), al- though the intensity of hve plant-tissue consumption re- mains unknown. It is highly likely that tissue-nonspecific oribatid mites that inhabited structural tissues, peat, and litter simply persisted into the Late Pennsylvanian, dur- ing which an essentially new, canopy-based, trophic level of primary consumers became estabhshed. While taxo- nomic membership of both trophic groups has changed through time, their basic ?cologie roles continue to the present day. Lineages of insect wood-borers occurring in modem gymnospermous and ai^ospermous trees are frequently documented as originating during the Triassic, However, this early Mesozoic origin for insects boring into above- ground hve and dead tissues in trees may be more appar- ent than real. The absence of any earlier damage by poten- tial Permian insect-borers may reflect taphonomic bias and absence of searchir^, particularly since penetrating hard tissues is a life habit that is plesiomorphic in blattoid, isopteran, and many holometabolous insect lineages {Hamilton, 1978; Crowson, 1981; Gepp, 1984; Nalepa, 1994), most which appear as body fossils during the Perm- ian and Triassic, While insect borer damage has not been described from Permian plants, by Late Triassic times ev- idently there was consumption of Hve tissue in standing woody plants. The incorporation of above-ground tis- sues?especially unaltered heartwood and live cambium and phloem?during the Early Mesozoic contributed new standit^-crop tissue to animal food webs. For nutritional- ly unrewarding heartwood, the expansion of wood-boring insects was undoubtedly associated with mycopha^ of fungi saprophytic on wood (Hickin, 1975; Crowson, 1981), This addition of new tissues previously exempt to borer consumption further integrated and sohdified links be- tween primary producers and arthropod consumers. CONCLUSIONS In this paper we have estabhshed major ecological pat- terns defining the fossil record of oribatid mites and the vascular plants they consumed. We have determined from available fossil occurrences and modem ecological studies some of the ?cologie roles that oribatid mites served in the degradation of plant tissues in Paleozoic coal swamps. From both our primary study of Euramerican coal-baU flo- ras and our review of the available literature on fossil ar- thropod plant-borers, we conclude the foUowit^ four points. (1) There was widespread consumption of diverse vascu- lar plant tissues in Late Carboniferous (Pennsylva- nian) Euramerican coal swamps by oribatid mites, producing spheroidal and ovoidal to cylindrical cop- rolites in the 45 jjtm to 110 jan size range, and tunnels within plant tissues that independently ranged from 100 Jim to 450 ]i,m. in diameter. (2) The known foss? history of oribatid mites is character- ized by both body-fossil and trace-fossil records. The body-fossil record commence during the Middle De- vonian but does not resume unt? the Jurassic, during which essentially modem taxa are encountered. By contrast, the trace-fossil record is largely confined to the Carboniferous, and provides iinique insights into feeding ecology of oribatid mites trom. Euramerican coal-swamp environments. These two fossU records are not only geochronologically complementary, but additionally reveal very different aspects of fossil ori- batid mite biology. Notably, the trace-fossil record opens a unique 75-m?llion-year window into oribatid mite life-history and behavior that closed during the Early Permian and did not reopen until the Quater- nary, (3) In seven major Euramerican coal-ball ?oras of Peim- sylvanian age, virtually all permineralized tissue types from the five dominant plant groups?lycopsids, calamites, fems, seed ferns, and cordaites?eidnbit tmmelingby oribatid mites. Hard and often minimal- ly altered postmortem tissues such as bark, fibrovas- cular bundles, and especially wood were bored, as were softer tissues, including various stem and fohar parenchyma and the gametophytic tissue of large seeds. Evidence for coprophagy of macroarthropod coprolites also is present. Wh?e mite-mediated detri- tivory is the most obvious form of arthropod consump- 344 LABANDEIRA ET AL. tion of plant tissues during the Early and Middle Pennsylvanian, insect herbivory assumes greater doc- umented importance diiring the Late Pennsylvanian, Lcnmediately after a major ?oral turnover from lycop- sid-dominated to tree fem-dominated coal-swamp communities. (4) Whereas a qualitatively diverse assemblage of above- ground, canopy-focused herbivores coexisted with mite detritivores during the Late Pennsylvanian, true herbivory of wood and other hard tissues in arbores- cent plants is not convincingly demonstrated until the Late Triassic, These and subsequent herbivore cul- prits were insect lineages feedings on cambium and phloem, and increasingly included mycophagy in the heartwood of gymnospermous trees and later of woody angiosperms. Meanwhile, the trace-fossil rec- ord of oribatid mite detritivory is absent for this time interval. Modem studies of htter decomposition in north-temperate forests indicate that their ground- level and below-ground trophic web of Pennsylvanian detritivory has persisted to the present, ACKNOWLEDGMENTS This work would not have been possible without the ef- forts of many colleagues who have retrieved, processed and curated the coal ball collection at the University of Il- linois for the past 35 years. Finnegan Marsh deftly drafted Fig. 1 and produced the layout for Figs. 2 to 12. We appre- ciate the conunentary of WA, DiMichele who reviewed an earlier draft of this manuscript. We acknowledge David Grimaldi of the Entomology Department at the American Museum of Natural History, Eandi Hansen of the Insti- tute of Ecology at the University of Georgia, David Head- rick of the Department of Entomology at the University of California, and Francis Hueber of the Paleobiology De- partment at the National Museum of Natural History for providir^ comments from unpublished data. 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DEVONIAN Givettian Devonacai?dae Padther Mtn. Fm., Gil- boa no Protochthoniidse Panther Mtn. Fm., G?- boa JURASSIC SiEemurian no Kydroietidu PankaipFm. Tithonian possibly Achipteriidae Eures Basin no Astegistidae Burea Basin no Trhypochthoniidae Burea Basin no Cymbaeremaeidae Burea Basin no Trhypoehthoniidae Burea Basin CRETACEOUS Santonian no Csin?iidae TaimjT amber no Plateremasidae Taimyr amber Campanian no GymnodainBeidae Manitaban amber no Oribatulid?? Manitoban amber TER?IAHY Danian possibly Oribatulidxe Sakalin amber Thaoetiaii possibly Hy?metidae Paskapoo Pm, Priabonian possibly Phthiraearidae Baltic amber yes Autognetidae Baltic amber no BrachychthoD?dae Baltic amber no Caleremaeida? Baltic amber no Can?s?dac Baltic amber no Carabodidae Baltic amber yes Carabodidae Baltic amber yes Carabodidae Baltic amber yes Caiaba?iiae Baltic an^r yes Cepheidaa Baltic amber possibly Baltic a??^^er no Chamobatidae Baltic amber DO Ast?gi&tidae Baltic amber no Baltic amber no Cymbaeremaeidae Baltic amber no Dadacidae Baltic as?)er no unassign??? Baltic amber no Baltic amber yes Er?rcaeidae Baltic amber no Pbanopelopida? Baltic amber no References Dtainaeanis selinkki Norton 19&S ProiJXhth?ttus g?boa Norton 1988 Mydrowtes sp. Acfiipteris (?) obscura Krivoltitsky 1976 Culioritiulajurasska Khvolutaky 1976 JuraiariiS s?mt?us Khvolutsky 1976 Jureretnus /oueo?a?u^ Kiivolutsky 1976 BaJaeochtli?T??s krasiiovi Krivolutsky 1976 Eocamisia sukatshafoi Bulanova-Zachvatina 1974 Bo?nit?tyn???o punctula?a Krivolutsfcy 1976 Genus and species unidentified Genus and ap?eles unidentified Soc?ia?inella th?r?AirU Eyab?nin 1976 Hydrozetes sp. Atropacarus midtipt?tu?tqlum (Sellnick 1919) Aaiogneta hJigHamellaium (Michael 1885) Brachyck?h?n???s sp. Caiersmaeus gleso Sellnick 1931 Camisia segnis (Hermann l?04) Carabodes coriacfus Koch 1836 Caraboades dissctms Selinick 1931 Carabodts gerben SeUnick 1931 Carabodes labyrinikicus (Michael 1879) Ctfiheus impUeaius (Sellnick 1919) Ceraioppi^ bipilts (Hermann 1804) Cham?bales diffi?lk Sellnick 1931 CuUnribula ?mita Sellnick 1931 Culteribuia saperba Sellnick 1931 "Cymbceremaeus" a?umin?ttus Si:\\nic)<: 1931 Darwtus (?) geftaderisis Sellnick 1931 Embfjtoiarus pergatui Sellnick 1919 EpBribaliila peltiaida Sellnick 1931 EremaetiS obhngus Koch 1S36 Eup?ops p?ttt?^?tlatus (Sellnick 1931) Norton et al., 1988, 1989 Nortonet al., 19S8, 1989 Sivhed & Walvork, 1978: Krivolutsky & Druk, 1986 Krivotutsky & KrassUov, 1977; Krivolutsky et al, 1990 Krivolutsky & Krassilov, 1977; Krivolutsky et al, 1990 Krivolutsky & Krassilov, 1977; Krivolutsky et al, 1990 Krivolutsky & Krassilov, 1977; Krivolutsky et al, 1990 Krivolutsky & Krassilov, 1977; Krivolutsky et aL, 1990 Zherikin & Sukacheva, 1973; BuUnsva-Zachvatina, 19T4; Keil< bach, 1982 Zherikin & Sukacheva, 1973; Bulamma-Zachvatina, 1974; Kri- volutsky & Ryabinin, 1976 Ewing, 1937; McAlpine & Mai?a, 1969; Krivolutsky ? Dnik, 1986 Ewing, 1937; McAlpine & Martin, 1969; Krivolutsky & DiuJi, 1986 Krividutaky & Ryabinin, 1976; Kiivollitslv A Omk, 1986 Baker & Wighton, 1984 Selkjck, 1919; Kellbach. 1962; Krivolutsky et al., 1990; Niedba- la, 1992 Michael, 1865; Krivolutsky et al., 1990 SeUnick, 1931; Keilbach, 1982; Krivolutsky et al, 1990 Sellnick, 1931; Keilbach, 1962; Krivolutsky et al., 1990 Karscb, 1884; Sellnick, 1919. 1931 sub Nothrus htrridus fir. Comisi? h?rridus h. fossilu?\ Coi?o? 1993 SeUnick, 1931; Kellbach, 1962; Krivolutsky et al., 1990 Sellnick, 1931; Keilbach, 1982; Krivolutsky et al., 1990 Michael Se George, 1879; Keilbach, 1982; Sellnick, 1931; Krivo- lutskyet al., 1990 SeUnick, 1931; Keilbach, 1962; Krivolutsky et al, 1990 Sellnick, 1919,1931; Keilbach, 1982; Krivolutsky et al., 1990 Sellnick, 1919; Keilbach, 1962; Krivolutsky & Dnik, 1936 SeUnick, 1931; Keilbach, 1962; Krivolutsky et al, 1990 SeUnick, 1931; KeUbach, 1982 SeUnick, 1931; Keilbach, 1982 SeUnick, 1931; Keilbach, 1962; Krivolutsky et al, 1990 SeUnick, 1931; Keilbach, 1962; Seiden, 1993a SeUnick, 1919; Krivolutsky et al? 1990 SeUnirfc, 1931; Keilbach, 1982; Krivolutsky et al? 1990 SeUni<^, 1919; Keilbach, 1932; Krivolutsky et al, 1990 SeUnick, 1931: Keilbach, 1962; Krivolutsky et al., 1990: Seiden, 1993a 352 LABANDEIRA ET AL APPENDIX 1. Continued. Taxon Family Deposit Endopha- gous taxon? References GaluntFta clcuvia Sellnick 1931 Gatumna diuersix S^Unick 1931 GradydorsuTTt asper SeUiuck 1919 Gymnodamoeus sepatisiis Selinick 1919 HermanntelUi C?Jt?iamerata Sel^ck 1919 Hermannie?ki tubert?data Sel?nicls 1919 ??cneremaetis fritschi S?llmck 1931 Liebstadia similifbrmis Sellui?k 1931 Liodes Q?iadrisc?iiaius (SeUnick 1919) Lvcoppia (?) simplex SeLlnick 1931 Melanosetes fodercUus Sellmck 19S1 Me?anosetes mallia?Jiu?S (Koch 1340J Micreremus reticuialus Sellnick 1931 Micreremus strobiculatus SeLLnick 1931 Neoribales orusstcus Selloick 1931 "?^otaspis" sp- N?ihrus it?autiis SeUuick 1919 "Ncthrus^ kiih? Katsch 1SS4 *'NQthrus" punctulum Karsch 1884 Odojtio?iph?us (?) sp. Oppia artgusium (S^llnick 1931) Oppia iiiTuiC?rna (SelLiuck 1919) Oppia medium (Sellnick 1931) Oppia sudnum ^Selloick 1931) Oribatella. mirttbiiis Sellnick 1931 Orib?iriiia pyr?pus (SeUnick 1919) Oribctritia trun^iucida (Sellnick 1931) Oripodct b?ltica S?Uiuck 1931 Otacepkius niger Sellnick 1931 Otacepheus pro^ignis SeUoick 1931 P?ateg?ocranus sidcaius (Karfich 1SS4) Fiatylicdes ensigertis (Sellnick 1919) Pr?t?ribaies langipUis Se?lnick 1931 Panci?riba?es sp. ScapkeremaetiS utidosus (SeUaick 1919) Scbeloriboies apertus Sellnick 1931 Scketoribaies ancoius SeUnick 1931 Scheioribctes setutit? S?lLnick 1931 Scuioribates peromalus Sellnick 1919 SpkcKnjz^tes conixxalus (Koch & Berendt 1854) Sph?er?zeies primus Selinick 1931 Strieremaeus cordifarmata^ Sellnick 1919 Sirkrernaeus iUibatus Sellnick 1919 Siii^t?beibe?Ia su-btrigona (Oud^msns 1900) l^tocepheus ?imilis SeUnick 1931 lixtxKymba nra Sellnick 1919 Trhypochihonius comiculaitis Sellnick 1931 Trhypochthonius bluiiiformis Sellnick 1931 Undu?oribates parvas (ScHnick 1931) Xenit?us tegeocraniformis (Sellnkk 1919) Marcoipeda rm^atianes P?rft? Alhnothrus sp. Galixmnatidae Galiunnatida? Eretn?eidae Gyi?madaiDaeidBe Hermanniellidae Hermanniellidae LJcneremandae Oribatulids? Liodid?? Oribatulidae C?ratozetidae CeratDzetida? MicreTemidae Micreremidae ParakaluiiQ matidae AdiipUnidae Nothridae Nothuidae Nothhdae Carabodidae Oppiidae Opp?dae Opp?dae Oppiidae OribaUllidae Oribc?titiidae Ofibotitiidae Ohpodidac Otocephe?dae Otocepheidae unas^ign^ Liodidae Haploietida? Myc?batidae Cymb?eremaeidae Scheloribatidae Scbe?Dribatida? Scheloribatidae unassigned Orstfit^tidae CeratO'Eetidae ?remaeldfle Suct?belb?dae Tectocepheldae Cymbaeremaeidae Trhypochthchiuid ae TrhypocKtb(}iuidae Unduloribatida? Xenillidae unassigned TrhypocKthonildae Scutovertiddae Baltic amber Baltic amber Baltiq amber Baititr amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amb?T Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amb&r Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Baltic amber Eocene Unspecified Zcna Oiauconitica Chattian El Mamey Fm.^ Domini- can amber El Mamey Fm., Domini- can amber possibly poASibly no yes yes no poaaibly no possibly yes yes possibly probably probably poEsiWy possibly possibly possibly Sellnick, 1931- Keilbach, 1982; Krivolutsky et al., 1990 Sellnick, 1931; Keilbach, 19?2; Krivolutsky et ai., 1990 Sellnick, 1919; Keilbach, 1982; Krivclutsky et ai,. 1990 Sellni?k, 1919; Ke?bach, 1982; Krivolutsky et al.. 1990 Sellnick, 1919; KeUbach, 1982; Kiivclutsky et ai., 1990 Sellnick, 1919; Keilbach, 1932; Krivolulaky et al, 1990 Sellntck. 1931; KeUbach, 1932; Krivolutsky et al.. 1990 Sellnick, 1931; Keilbach, 1982; Krivolutsky et al,. 1990 Sellnick, 1919; Keilbach^ 19B2; Krivolutsky et al,. 1990 Seiblick, 1931; Keilbach, 1982; Krivolutsky et al., 1990 Sellnick. 1931; Keilbach, 1962; Krivolutsky et al., 1990 Sellnick. 1931; Kellbach, 1982; Krivolutsky et al.. 1990 Sellnick. 1931; Keilbach, 1982 Sellnick, 1931; Keilbach, 1982; Krivolutsky et al,. 1990 Sellnick, 1931; Keilbach, 1982; Krivolutsky et al.. 1990 Seiblick. 1919; KeUbach, 19S2 Setinick. 1919; KeUbach. 1982; Krivolutsky et al., 1990; Seiden, 1993a Kaisch, 1894; Keilbach. 1932 Karscb, 1884; Keilbach, 1982 Sellnick, 1931; KeUbach, 1982; Seiden, 1993a Sellnick, 1931^ KeUbach, 1982; Krivolutsky et al., 1990 Sellnick, 1919,1931; Keilbach, 19S2; Krivolutsky et al., 1990 Sellnick. 1931; KeUbach, 1982; Krivolutsky et al., 1990 Sellnick^ 1931; Keilbach, 1982; Krivolutsky et al.. 1990 Sellnick, 1931; KeUbach, 1982; Krivolutsky et al? 1990 Sellnick. 1919,1931: Keilbach, 1982; Krivolut?ky et al., 1990 Se?nick, 1931; KeUbach, 1982; Krivolutsky et al.^ 1990 Sellnick, 1931; Keilbach, 1982; Krivolutsky et al., 1990 SeUnick, 1931; KeUbach, 1982; Krivolutsky et al., 1990 Sellnick, 1931; KeUbach. 1932; Krivolutsky et aL, 1990 Karsch, 1834; Sellnick, 1919; KeUbach, 19B2; Krivolutsky & Druk, 1936 Sellnick, 1919,1931; Ke?bach. 1982; Krivolutsky & Dnik, 1986; Knvolutsl?y et al., 1990 Sellnick, 1931; KeUbach, 1982; Krivolutsky et al., 1990 Seilnick, 1931; KeUbach, 1982; Krivolutsky et al., 1990 Selhiiek, 1931; KeUbach, 1932; Krivolutsky et aL, 1990 Sellnick 1919, 1931; Keilbach, 1952; Krivolutsky et al., 1990 Sellnick, 1931; Keilbacb, 1982; Krivolutsky et al., 1990 SeUnick, 1931; KeUbach, 1992; Krivolutsky et al., 1990 Sellnick, 1919; KeUbach, 1932; Krivolutsky & Druk, 1980; Sellnick, 1919; KeUbach, 1982; Krivolutsky & Ortik, 1986; Kri- volgtsky et aL, 1990 Sellnick, 1931; K?ilbach, 1932; Krivolutsky et al., 1990 Sellnick, 1919; Keilbacb. 1982 Sellnick, 1919; KeUbach, 1982; Krivolutsky SL Druk, 1936- Kri- volutsky et al-, 1990 SeUnick, 1931; Keilbach, 1982; Oudemans, 1900; Seiden, 1993? Sellnick, 1931; Keilbacb, 1932; Krivolutsky et al., 1990 Sellnick, 1919. 1931; KeUbach, 1982; Krivolutsky et al., 1990 SeUnick, 1931; Keilbach, 1982; Krivolutsky et al., 1990 SeUnick, 1931; KeUbach^ 1932; Krivolutsky et al., 1990 Sellnick, 1931; KeUbach. 1932; Krivolutsky et al.. 1990 SeUnick, 1919; Keilbach. 1982; Krivolutsky et al.. 1990 P?rei, 1938 Norton & Poinar, 1993 Norton & Poinar, 1993 ORIBATID MITES IN COAL-SWAMP FORESTS 353 APPENDIX 1. Continued. Taxon Family Deposit Endopha- gous taxon? References Klapparic/as sp. DtAicheremmus sp, Liodes sp. ^Oppia sp, Orib?triti? sp. OripodcL sp. Soccuiobaiea sp. T?leiotiodes sp. genas and species uiudenti?cd Arthrwrriex kurdi {W?&lley 1971) Bfftoifca?f?r chiajKtssnsis (Woolley 1971) Liodes bre?iiarsus (W?otley 1971) ^?ochhr?bo?v?a smiiki tWoolley 1971) Opp?a sctifer {y?w?ey 1971) Oppia maicarut tWoollcy 1971) Faropirn^us denaius (Woolley 1971) Sckeloribauts durhami (Woolley 1971) Betbites disodotia Pampaloni 1902 Carabodiies paaesii Pampakimi 1902 Oppttes raeiilii Pft?npalcuu 1902 Crot0n? Chiapas amber Simojovel F?n,, Chiapas amber SerravaUian Sicilian amber Sicilian amber Sicilian amber Zanclian Australian amber; Al^ lendale, Victoria Piacenzian pt?ssibly possibly Beaufort Fm,; Prince no Patrick Island, North West Territoriea no QUATERNARY Numerous occurrences; see Elias (1994) Norton & Poinar, 1993> sub Carabodes; R. M. Reeves, pers. comm. Norton & Foinar. 1993 Norton & Pt^inar, 1993 Norton & Po?nar, 1593 Norton & Poinar. 199S Norton & Poinar, 1993 Norton & Poinai, 1993 Norton St Poinar> 1993 Norton & PoiJiart 1993 Norton St Poinar, 1993 Norton & Poinar, 1993 WooUey. 1971; Keilbach, 1982; Norton & Poinar, 1993 Woolley, 1971-, Keilbach, 1982: Norton & Poinar, 1993 Woolley. 1971; Keilbach, 1982; Norton 6t Peinar, 1993 WooUey, 1971; KoUbaeh. 1982; Norton & Poinar, 1993 Woolley, 1971; KeilbacK^ 1982; Norton & Poinar, 1993 Woolley. 1971; KeQbach, 19S2; Norton & PE^inar, 1993 WooUey, 1971; Keiibach, 1982; Norton & Poinar, 1993 WooUey, 1971; KcUbach. 1982: Norton & Panar, 1993 Pampaloni, 1902; Krivolut&ky et al., 1990 Pampal?ni, 1902; Krivolutsky et al.. 193? Pampaloni, 1902; Krivolutsky et aln. 1990 Wometsley. 1956; Keilbach. 19S2 Gosolova et al.. 19?5; Krivolutslcy & Druk, 1986 Gosolova et aL, 1985; Krivolutsky & Druk, 19S6 Gosolova et al., 1965; Krivolutsky S? Dmk. 1986 Gosolova et al,, 1985; Krivolutsky & Dmk, 19S6 Gosolova et al., 1985; Krivolutsky & Druk, 1936 Gosolova et al., 1985; Krivolutsky & Dmk, 1936 Gosolova et al., 1985; Krivolutsky & Druk, 19S? Matthews ^ Ovenden. 1990; Behan-Pelletier & Ryabinin. 1991 Druk, 1982; Krivolutsky et al., 1990