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Ichnos

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Taxonomic reassessment of the ichnogenus beaconichnus and additional examples from the Carboniferous of Kansas, U.S.A. Luis A. Buatois a; M. Gabriela Mángano a; Christopher G. Maples a; William P. Lanier b a Kansas Geological Survey, The University of Kansas, Lawrence, KS, USA b Department of Earth Sciences, Emporia State University, Emporta, KS, U.S.A.

To cite this Article Buatois, Luis A., Mángano, M. Gabriela, Maples, Christopher G. and Lanier, William P.'Taxonomic

reassessment of the ichnogenus beaconichnus and additional examples from the Carboniferous of Kansas, U.S.A.', Ichnos, 5: 4, 287 — 302 To link to this Article: DOI: 10.1080/10420949809386427 URL: http://dx.doi.org/10.1080/10420949809386427

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Ichnos, Vol. 5, pp. 287-302,1998 Reprints available directly from the publisher Photocopying permitted by license only

Taxonomic Reassessment of the Ichnogenus Beaconichnus and Additional Examples from the Carboniferous of Kansas, U.S.A. LUIS A. BUATOISa,*, M. GABRIELA MÁNGANOa, CHRISTOPHER G. MAPLESa and WILLIAM P. LANIERb,† a

Kansas Geological Survey, 1930 Constant Avenue, Campus West, The University of Kansas, Lawrence, KS 66047, USA;b Department of Earth Sciences, Emporia State University Emporta, KS 66801, U.S.A.

The ichnogenus Beaconichnus (Gevers 1973), an arthropod trace fossil, includes very different forms that comprise five ichnospecies, namely B. darwinunt (Gevers 1971), B. gouldi (Gevers 1971), B. ahtarcticum (Gevers 1971), B. giganteum Gevers and Twomey 1982, and B. wrrighti Gevers and Twomey 1982. The original diagnosis of Beaconichnus is rather vague and potentially may accomodate virtually every arthropod trackway described from the fossil record. In view of these problems, the validity of Beaconichnus is reassessed and each of its ichnospecies is reviewed. We conclude that B. darwinum is a junior synonym of Diplopodichnus biformis Brady 1947; B. antarcticum should be regarded as Palmichniunt antarcticum; and B. wrighti is a nomen nudum. Additionally, we agree with previous proposals in considering B. gouldi as the senior synonym of B. giganteum, and including it in Diplichnites Dawson 1873. Therefore, we suggest that the ichnogenus Beaconichnus is best disregarded. Additionally, we describe specimens collected from the Late Carboniferous Tonganoxie Sandstone Member (Stranger Formation) of eastern Kansas, ascribed herein to Diplopodichnus biformis and Diplichnites gouldi, which include examples of intergradations between both ichnotaxa, and provide synonymy lists for both ichnospecies. Keywords: Arthropod Trace Fossils, Ichnotaxonomy, Carboniferous, Kansas * Corresponding author. † Deceased. 287

INTRODUCTION Arthropod trackways are among the most complex groups of traces from an ichnotaxonomic viewpoint. Outstanding problems include: a remarkable variability of trackways within single specimens or populations, strong preservational constraints on trackway morphology, an absence of a broad consensus on the hierarchical significance of ichnotaxobases and an inadequate diagnosis of several ichnogenera and ichnospecies. The taxonomic problems of this group are particularly significant because arthropod trackways are important components of many trace fossil assemblages, especially in Paleozoic nonmarine environments (e.g. Brady, 1947; Walter, 1983; Walker, 1985; Demathieu et al, 1992; Acenolaza and Buatois, 1993). Additionally, inconsistencies in taxonomic practices and formulation of new ichnotaxa on the basis of uncertain morphological features and fragmentary material have resulted in a plethora of ichnotaxa, some of them undoubtedly junior synonyms of formerly proposed forms. As noted by Buatois et al. (1995), this type of taxonomic problem represents a serious bias for the analysis of ichnodi-

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versity trends of nonmarine ichnofaunas through time (c/. Buatois and Mângano, 1993). Recently, Trewin (1994) attempted to solve part of these difficulties by providing a preliminary system for trackway description and identification, establishing a set of trackway attributes or ichnotaxobases, which may prove to be useful in the analysis of new ichnotaxa and the revision of formerly established trace fossils. Similarly, Braddy (1995a) clarified trackway terminology, listed a series of diagnostic characters of arthropod trackways, and suggested that only morphological features indicative of particular tracemakers should be used in ichnospecific diagnoses. There is still, however, disagreement about which characters are significant at ichnogeneric and ichnospecific ranks. The ichnogenus Beaconichnus (Gevers 1973), the subject of this contribution, is a microcosm of the problems previously outlined. Its original diagnosis is so broad that it may include most of the arthropod trackways known from the fossil record. The situation is further complicated by the fact that remarkably different morphological patterns were included within the ichnogenus as different ichnospecies, namely B. darwinum (Gevers 1971), B. gouldi (Gevers 1971), B. antarcticum (Gevers 1971), B. giganteum Gevers and Twomey 1982, and B. xvrighti Gevers and Twomey 1982. We discuss herein the accommodation of all Beaconichnus ichnospecies within other ichnogenera, such as Diplichnites, Palmichnium, and Diplopodichnus, and conclude that Beaconichnus is a poorly-defined ichnotaxon, recommending its abandonment. As part of a broader study on the ichnology of the Late Carboniferous Tonganoxie Sandstone Member (Stranger Formation) of eastern Kansas, specimens ascribed herein to Diplopodichnus biformis Brady 1947 and Diplichnites gouldi (Gevers 1971) were collected. These specimens provide additional evidence for an ichnotaxonomic reassessment of Beaconichnus. The purpose of this contribution is to discuss the taxonomy of the ichnogenus Beaconichnus, to

provide evidence for the reassessment of its different ichnospecies and to describe this new material.

ICHNOTAXONOMIC REASSESSMENT OF BEACONICHNUS The ichnotaxonomic status and history of Beaconichnus are remarkably confused. Originally defined by Gevers (in Gevers et al., 1971) as Arthropodichnus, it subsequently was emended to Beaconichnus by Gevers (1973) due to preoccupation of the name (Arthropodichnus Chiplonkar and Badve 1970). One of the main problems with the ichnogenus Beaconichnus lies in its extremely vague diagnosis. Gevers et al. (1971, p. 92-93) characterized it as "Epistratal trails made by arthropods... Trails display evidence of symmetrically distributed rows of walking appendages on either side of mid-line. Footprints and telson mark not necessarily distinguishable." According to this diagnosis, many well-established arthropod trackway ichnotaxa could be included in Beaconichnus (e.g. Palmichnium, Umfolozia, Maculichna). Gevers et al. (1971) proposed three ichnospecies: B. darwinum, B. gouldi, and B. antarcticum (Table I). B. darwinum was regarded as consisting of two parallel grooves, which may or may not display foot imprints. B. gouldi was described as "paired parallel rows of closely spaced foot imprints, forming grooves in some cases." B. antarcticum was considered as characterized by the presence of median telson marks, from which diverge rows of imprints. Differences in the size of each ichnospecies were also emphasized. B. darwinum ranged in external width from 9 to 18 mm, B. goüldi was 22-43 mm wide, and B. antarcticum varied from 290 to 300 mm wide. Subsequently, Häntzschel (1975) listed and figured a fourth ichnospecies as Beaconichnus giganteum Gevers, not included in the original paper. This ichnospecies, however, was formally defined later by Gevers and Twomey (1982) without any

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mention to HMntzschel's comment. B. giganteum was considered as consisting of large trails, 95200 mm wide, with two rows of imprints arranged in clusters of two to four. Gevers and Twomey (1982) also proposed a fifth ichnospecies, B. wrighti, which was regarded as characterized by the existence of chevron-shaped imprints aligned in two parallel rows. This latter ichnospecies, however, was not figured and neither a formal diagnosis nor type material were provided. Therefore, B. wrighti should be regarded as a nomen nudum (Table I). Bradshaw (1981), however, demonstrated a complete gradation in size from B. darwinum to B. gouldi to B. giganteum and suggested that there was no basis for maintaining B. giganteum as a separate ichnospecies, which was regarded as a synonym of B. gouldi. Additionally, Bradshaw (1981) ascribed B. gouldi to the ichnogenus Diplichnites, a taxonomic proposal accepted by Johnson et al. (1994) and endorsed in this contribution (Table I) (Fig. 1A). Bradshaw (1981) stressed also that B. antarcticum shares with Paleohelcura the linear oblique arrangement of the imprints and considered the larger size of the former as the only difference between both ichnotaxa. Consequently, she suggested to place B. antarcticum as an ichnospecies

^

^

A. Diplichites gouldi B. Palmichnium antarcticum C. Diplopodichnus biformis

FIGURE 1 Line drawings of ichnotaxa formerly included in Beaconichnus. (A) Diplichnites gouldi. (B) Palmichnium antarcticum. (C) Diplopodichnus biformis

of Paleohelcura, P. antarcticum. Subsequently,

Walker (1985) noted similarities between Paleohelcura and the ichnogenus Stiaria Smith 1909, pointing out that the latter, however, exhibits a more variable number of imprints and patterns. She placed the material illustrated by Brady (1947, Fig. 1.4) as P. tridactyla in Stiaria quadripe-

dia. This decision was recently criticised by Trewin and McNamara (1995), who stated that the median groove, apparently essential to the diagnosis of Stiaria, is not necessarily present in Paleohelcura. We believe that, at least provision-

TABLEI Summary of Beaconichnus ichnotaxonomy Gevers et al. (1971) Arthropodichnus darwinum

Gevers (1973)

Gevers & Twomey (1982)

Bradshaw (1981)

Beaconichnus darwinum

Arthropodichnus darwinum (partim)

Beaconichnus gouldi (partim)

Arthropodichnus gouldi (partim)

Beaconichnus gouldi (partim)

Arthropodichnus antarcticum

Beaconichnus antarcticum

Beaconichnus gouldi

Diplichnites gouldi

Keighley & Pickerill (1996)

This paper

Diplopodichnus biformis

Diplopodichnus biformis Diplichnites gouldi Diplopodichnus biformis

Beaconichnus giganteum Beaconichnus zvrighti

Paleohelcura antarcticum

Palmichnium antarcticum

Diplichnites gouldi

Diplichnites gouldi nomen nudum

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ally, both Stiaria and Paleohelcura should be retained as different ichnotaxa. Five ichnospecies of Paleohelcura are known: P. tridactyla Gilmore 1926, P.? lyonsensis Toepelman and Rodeck 1936, P. dunbari Brady 1961, P. delicatula Fischer 1978, and P. badensis Kozur et al. 1994. Mesichnium benjamini Gilmore 1926 has been regarded as a variation of Paleohelcura and may therefore be considered as a sixth ichnospecies, P. benjamini (Kozur et al, 1994; Braddy, 1995a). Fischer (1978, p.25) stated that "P. delicatula differs from P. tridactyla in its smaller size, in having 'comma-shaped dactyls', and in showing a tendency towards a more opposite alignment of sets of tracks". However, he did not address the distinction between P. delicatula and the remaining Paleohelcura ichnospecies. The shape of imprints is not dear in the illustration (Fischer, 1978; fig. 9a) and the smaller size is probably of little value in this case as a diagnostic criterion (Pickerill, 1994). However, the opposite symmetry may be useful to distinguish P. delicatula from the other Paleohelcura ichnospecies. Braddy (1995a) stated that P. badensis only differs from P. tridactyla by its smaller size, and accordingly regarded it as its junior synonym. Additionally, Braddy (1995a) also considered P. dunbari and P. lyonsensis as junior synonyms of P. tridactyla, although both ichnospecies display distinctive imprint arrangement. He suggested that these ichnospecies record behavioral variations of the P. tridactyla pattern, with P. dunbari representing faster locomotion and P. lyonsensis reflecting slower movements. However, it could be argued that both ichnospecies should be considered valid because ichnotaxa are currently based on morphology as an expression of animal behavior (Bromley, 1990). None of the Paleohelcura ichnospecies, however, exhibit the external width and imprint size of B. antarcticum. Additionally, B. antarcticum displays staggered symmetry. Accordingly, we suggest that B. antarcticum should not be included in Paleohelcura (Table I) (Fig. IB).

Similar to the ichnogenus Palmichnium Richter 1954, Beaconichnus antarcticum consists of large symmetrical trackways that displays two rows of imprints, which occur in groups of three and are separated by a central continuous mark. Similarities between both ichnotaxa formerly were noted by Gevers et al. (1971) and Häntzschel (1975). However, Gevers et al. (1971) stated that B. antarcticum differs from Palmichnium by the angles between the telson mark and the set of footprints and by their isolation from the telson mark. According to Gevers et al. (1971). B. antarcticum shows an angle of 35° between imprint groups and the telson mark, whereas P. palmatum displays an angle of 45° (Hanken and Stormer, 1975). Nevertheless, the diagnosis of Palmichnium, as emended by Briggs and Rolfe (1983), includes trackways with high angles between series and the axis, without further specifications. Regarding the isolation of imprints with respect to the telson mark as a distinctive character of B. antarcticum, this is true only with respect to P. palmatum, which displays imprints linked to the telson mark. Nevertheless, of the remaining four Palmichnium ichnospecies, P. stoermeri, P. kosinskiorum, and P. macdonaldi display imprints isolated from the central mark (Briggs and Rolfe, 1983; Braddy, 1995b) and P. pottsae lacks a medial impression entirely (Braddy and Anderson, 1996). According to Trewin and McNamara (1995), the diagnostic feature that distinguishes Beaconichnus antarcticum from Palmichnium is the presence of alternate symmetry instead of opposite symmetry, as displayed by the latter. However, symmetry in Palmichnium is variable, with P. palmatum, P. kosinkiorum and P. macdonaldi displaying opposite symmetry and P. stoermeri and P. pottsae having staggered symmetry (Richter, 1954; Hanken and Stornier, 1975; Briggs and Rolfe, 1983; Braddy, 1995b, Braddy and Anderson, 1996). Accordingly, there are no significant criteria that allow separation of B. antarcticum from Palmichnium, and therefore this ichnospecies is regarded as Palmichnium antarcticum. A forthcoming paper

BEACONICHNUS: CARBONIFEROUS OF KANSAS

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by Braddy and Millner (Braddy, written communication, 1996) discusses the B. antarcticum problem and refers this ichnospecies to Palmichnium.

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Distinctions between bilobate burrow ichnotaxa (e.g. Cruziana, Didymaulichnus) and paired trails (e.g. Diplopodichnus) and related terminology were discussed in detail by Keighley and The remaining ichnospecies, B. darwinum; conPickerill (1996), so there is no need of repetition sists of two parallel epichnial grooves. According here. Fedonkin and Palij (in Palij et al., 1979) creto Gevers et al. (1971), foot imprints may or may ated the ichnogenus Bilinichnus (with its single not be discernible. Similarly, the ichnogenus Diploichnospecies B. simplex) to name parallel narrow podichnus, erected by Brady (1947) from the eolian grooves previously described in open nomenclaCoconino Sandstone (Permian) of Arizona, is charture by Fedonkin (1977) from the Vëndian of acterized by two or three parallel grooves. This northwest Russia. Keighley and Pickerill (1996) author also stated that occasionally foot imprespointed out the problems of explaining the mode sions are visible in the two outer grooves. Speciof formation of these structures prior to the mens of Diplopodichnus described by Brady (1947) dawn of hard part-bearing metazoans. These are smaller than the type specimen of Beaconichnus authors regarded Bilinichnus as a nomen dubium, darwinum (4-6 mm wide versus 9-18 mm wide but provisionally retained it and differentiated it respectively). However, as stated before, size is not from Diplopodichnus by the presence of striate or currently considered a significant ichnotaxobase punctuate ornamentation in the furrows of the (Pickerill, 1994). Morphologically, the two-grooved latter. However, imprints are not necessarily prespecimens of Diplopodichnus are indistinguishable served in Diplopodichnus, and Bilinichnus may from B. darwinum. Accordingly, we consider B. dartherefore be considered as its potential junior winum as a junior synonym of Diplopodichnus and synonym. We have tentatively synonymised suggest its inclusion under D. bijbrmis, in a single occurrences of B. simplex until its holotype be ichnospecies (Table I) (Fig. 1C), a conclusion also reexamined. reached independently by Keighley and Pickerill Although originally proposed for non-trilobite (1996). Neoichnologic experiments with myriatrackways (Dawson, 1873), the ichnogenus pods, conducted by Brady (1947), showed that Diplichnites was adopted later for trilobite locotwo-grooved trails were formed while the animal motion traces by Seilacher (1955). This ichnogewas moving upslope, whereas the three-grooved nus currently is used for simple, biserial traces recorded downslope displacements. Theretrackways. According to its original description fore, the presence of the third groove is not (Gevers in Gevers et al., 1971), Diplichnites gouldi regarded as a significant ichnotaxobase, and we is difficult to differentiate from Diplopodichnus suggest that Diplopodichnus biformis should include biformis. Gevers et al. (1971) argued that Diplopodboth two- and three-groove forms in this case. ichnus biformis (their Beaconichnus darwinum) is We also have included in Diplopodichnus biformis characterized by two parallel grooves that show small imprints, but also recognized that imprints another taphonomic variant that consists of furmay not be discernible in some cases. Additionrows preserved in positive hyporelief that may disally, they stated that Diplichnites gouldi consists of play adjacent, tiny, appendage imprints. This latter case records the preservation of Diplopodichnus closely spaced imprints forming two parallel biformis on bed soles, instead of bed tops. Append- rows that may form grooves in some cases. Clearly, both ichnospecies overlap morphologiage imprints of this preservational variation of D. cally. The suggestion that Diplichnites gouldi is biformis resemble lateral markings exhibited by Cochlichnus antarcHcus (Tasch, 1968). However, the wider than Diplopodichnus biformb was rejected nature of the lateral markings in C. antarcticus is by Bradshaw (1981), who demonstrated a complete size gradation between both ichnospecies. still unclear (cf. Fillion and Pickerill, 1990).

LUISA.BUATOISe«a/.

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FIGURE 2 Examples of Diplopodichnus biformis and Diplichnites gouldi from the Tonganoxie Sandstone at Buildex Quarry. (A) Enlargement of B showing D. gouldi intergrading with D. biformis (upper right), and D. gouldi trackway cross-cut by D. biformis burrow (center). Positive hyporelief. TB-9-95. (B) General view of specimens of D. biformis associated with D. gouldi. Note that D. biformis cross-cuts D. gouldi (center), and that the poorest preserved imprints occur in burrow segments that exhibit secondary successive branching, indicating formation of cummulative structures. Compound specimens are also displayed (upper center). Positive hyporelief, TB-9-95. (C) Diplopodichnus biformis consisting of two parallel ridges. Positive hyporelief. TB-8-96. (D) Diplichnites gouldi. Note the presence of unconnected, ellipsoidal imprints. Positive hyporelief. TB1-95. Scale bars = 1 cm

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FIGURE 3 Examples of Diplopodichnus biformis and Diplichnites gouldi irom the Tonganoxie Sandstone at Buildex Quarry. (A) Distinctive D. gouldi trackways become poorly defined and grade to the right into deformed traces of D. biformis (upper center). Note also the presence of discrete D. gouldi trackways (lower left) and D. biformis exhibiting secondary successive branching (right center). Negative epirelief. TB-3-96. (B) D. biformis preserved as two parallel, straight to sinuous epichnial grooves. Note sharp turns of direction. TB-5-96. (C) D. biformis that exhibits imprint markings towards the margin of the burrow. Positive hyporelief. Small grazing traces assodated. TB-4-96. Scale bars = 1 cm

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Furthermore, morphological transitions between

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Diplichnites gouldi and Diplopodichnus biformis

have been detected in the specimens from Buildex Quarry. It is not uncommon to observe specimens that laterally grade from one ichnospecies to the other (Figs. 2A-B, 3A). A similar situation recently has been documented by Johnson et al. (1994) from the Ordovician of the English Lake District. Accordingly, we consider both forms as end-members of a continum of transitional examples and restrict Diplichnites gouldi to those trackways that do not form continuous grooves. Therefore, those examples of Diplichnites gouldi that show grooves, as described by Gevers et al. (1971), should be included in Diplopodichnus biformis. The fact that certain- ichnotaxa may intergrade into another form has been analysed recently in detail by Pickerill (1994) and Pickerill and Narbonne (1995), who proposed the term "compound ichnotaxa" for such cases.

GEOLOGIC SETTING OF THE KANSAS SPECIMENS

The specimens documented herein were collected from the Late Carboniferous (Virgilian) Tonganoxie Sandstone Member of the Stranger Formation (Douglas Group) in Buildex Quarry, located southwest of Ottawa, Franklin County, Kansas (Fig. 4). The Tonganoxie Sandstone records part of the infill of a tide-dominated estuarine paleovalley (Larder et al, 1993; Archer et al, 1994; Feldman et al, 1995). The sedimentology of the Tonganoxie Sandstone at Buildex Quarry recently was analyzed in detail by Lanier et al (1993), who concluded that the Buildex succession was deposited on an upper tidal flat, near or at the fluvial-estuarine transition of the estuarine paleovalley. At this locality, the Tonganoxie Sandstone is 9 m thick and consists of very fine-grained sandstones, siltstones, and mudstones (Lanier et al., 1993).

LOCATION MAP NEBRASKA KANSAS

\

Buildex Quarry

l Douglas Group outcrop 50 mi 80 km

FIGURE 4 Location map of the Buildex Quarry (based on Lanier et al, 1993)

Trace fossils from the Buildex Quarry were first documented by Bändel (1967), who described and illustrated specimens attributed to isopods and limulids, leaving them in open nomenclature. The ichnologic content of the Tonganoxie Sandstone was subsequently mentioned in a series of sedimentologic and stratigraphic papers (e.g. Larder et al, 1993; Archer et al, 1994; Feldman et al, 1995; Tessier et al, 1995) and analyzed in detail by Buatois et al (1997). The specimens described in this contribution occur on the soles and tops of normally graded and parallel-laminated, coarse-grained, gray siltstone beds. These strata display remarkable lateral continuity and rhythmic stratification. The trace fossil-bearing deposits belong to the planar-bedded-and-laminated fades of Lanier et al. (1993).

TRACE FOSSIL TAPHONOMY

In the present case, the existence of compound ichnotaxa indicates that Diplopodichnus biformis

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and Diplichnites gouldi were produced by the same animal. However, no transitions have been observed between the smallest specimens of D. biformis and D. gouldi, suggesting that, in this particular case, they probably record activity of a different tracemaker. Intergradation between D. biformis and D. gouldi was also noted by Johnson et al. (1994), who found that both ichnotaxa are similar in size, display similar ethologic patterns, and commonly intergrade one into the other. In their case, Diplichnites consistently overcrossed Diplopodichnus where superimposed. Intergradations between D. biformis and D. gouldi also were mentioned by Trewin and McNamara (1995).

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(Fig. 3A). This explanation is consistent with the environmental setting proposed by Lanier et al. (1993), who suggested frequent and brief periods of subaerial exposure of the sediment with high water content during deposition. A potential ethologic explanation also deserves a brief comment. As the ichnogenus Cruziana records trilobite furrowing and related trilobite-produced Diplichnites represents walking activities, a similar situation may be assumed for transitions between Diplopodichnus biformis and Diplichnites gouldi. Such a possibility cannot be disregarded in the case of intergrading specimens. In other cases, both ichnotaxa are superimPreservation of Diplopodichnus or Diplichnites posed and, in contrast to the Lake District specimens, Diplopodichnus cross-cuts Diplichnites. may reflect either taphonomic processes or Moreover, the quality of preservation of the behavioral patterns. Johnson et al. (1994) discounted the possibility that Diplichnites actually imprints within Diplopodichnus specimens is highly variable. The poorest preserved imprints represents undertracks of a second generation of occur in furrow segments that exhibit secondary trackways formed after deposition of a thin successive branching and thus record the veneer of sediment on the Diplopodichnus horirepeated passage of the tracemaker within the zon because fine laminations necessary for the furrow (Figs. 2A-B and 3A). Additionally, these formation of undertracks were absent and both segments are deeper than the other segments of ichnotaxa intergraded within a single trace. They the trace. Therefore, we condude that in these also noted that the parallel lines of the trace were cases obliteration of imprints and formation of more widely spaced in certain parts of the trackcontinuous parallel elements, typical of Diplopodway due to local variability in sediment fluidity. ichnus, are related to the repetitive passage of the Additionally, experiments with the woodlouse animal through the same portion of the furrow. Oniscus asellus showed that grooves were proAccordingly, these furrows actually represent duced in wet substrates and trackways were cummulative structures. formed when the substrate was drier. Accordingly, Johnson et al. (1994) suggested that the preservational contrast records drying of the substrate, with Diplopodichnus formed in a wet SYSTEMATIC PALAEONTOLOGY substrate and Diplichnites in a drier sediment as animals traversed from one pool to another. Preservation of Diplichnites trackways in damp, Ichnogenus Diplopodichnus Brady 1947 firm sediment also was recorded by Wright et al. Type ichnospecies. Diplopodichnus biformis Brady, (1995) from the Silurian of Newfoundland. 1947 by monotypy. Keighley and Pickerill (1996) noted that Brady (1947) designated a slab with Detailed analysis of the Buildex specimens numerous traces as the holotype. Because only a suggests that this second alternative may explain single specimen can be nominated as a holotype, some of the observed transitions. In some specimens, distinctive Diplichnites trackways become Keighley and Pickerill (1996) proposed a lectotype from Brady's material. poorly defined and grade into deformed trails

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Emended diagnosis. Unbranched, straight to winding, horizontal trace consisting of two or three parallel epichnial grooves or hypichnial ridges that may display slightly impressed and closely-spaced elongate to ellipsoidal imprints perpendicular to the trace axis. Grooves/ridges separated by a distance equal or greater than the width of individual grooves/ridges. Imprints may also mark the margins of a furrow preserved as positive hyporelief (modified from Brady, 1947, and Keighley and Pickerill, 1996).

v?1978

Diplopodichnus bif ormis Brady 1947 Fig. 1C; Fig. 2A-C; Fig. 3A-C

71983

71979 71979 1981 71982 71983

1984 .1909 .1947 .1957

71957 .1967 .1971

.1971 .1971 .1971 .1973 .1975

71977

Tramway gutters Smith, p. 19-20, figs 31 and Fig. 32 Diplopodichnus biformis Brady, p. 469470 Isopodichnus osbornei Glaessner, p. 104-108, pi. 10, fig. 2; pi. 11, figs. 1 and 2 Isopodichnus osbornei Glaessner, p. 104-108, pi. 11, fig. 3 (partim) arthropod trails Rocha-Campos, fig. 42 Arthropodichnus darwinum Gevers et al., p. 93, pi. 19, figs. 1 and 2; figs. 3 and 4 (partim) Arthropodichnus gouldi Gevers et al, p. 93, pi. 19, figs. 3 and 4 (partim) Diplichnites sp. Savage, p. 225-226 (partim), fig. 9A Isopodichnus sp. Savage, p. 227-228, %• 13 Beaconichnus darwinum Gevers, p. 1002 Beaconichnus darwinum Häntzschel, p. W45, fig. 27.1c [copy of Gevers et al, 1971, pi. 19, fig. 1] and fig. 27.1b (partim) [copy of Gevers et al, 1971, pi. 19, fig. 3] parallel furrows Fedonkin, p. 184, pi. 2a

71984 71984

.1984 1985 71985 non 1985 non 1985 non 1986

71990 non 1990 v. 1991 v?1991

v.1993

Scolicia sp. Acefiolaza, p. 24, fig. 1-11.2 Bilinichnus simplex Pali) et al, pi. 61. 1-2 [copy Fedonkin, 1977, pi. 2a] Bilinichnus sp. Palij et al, pi. 53.6 Isopodichnus prdblematicus Holub and Kozur, p. 99, pi. 5.3 cf. Beaconichnus darwinum Pollard et fl/.,p.82. Bilinichnus simplex Palij et al, p. 87, pi. 60.1-2 [pi. 60.1 copy Fedonkin, 1977, pi. 2a] Bilinichnus sp. Palij et al, p. 87, pi. 53.6 [copy Palij et al, 1979, pi. 53.6] cf. Beaconichnus Pollard and Walker, p. 572. Cruziana sp. Pickerill et al, p. 420, fig.5B Scolicia isp. Fillion and Pickerill, p. 28, fig. 7c [copy Pickerill et al, 1984, fig.5B] Isopodichnus sp. Guerra-Sommer et al, 1984, p. 133, pi. 4, fig. 1 cf. Beaconichnus Walker, p. 295, fig. 9d double crawling trail Fedonkin, pi. 28.1 Bilinichnus simplex Fedonkin, p. 113, pi. 26.6 (= Didymaulichnus) Bilinichnus simplex Paczesna, p. 259, pi. 4.4 (= Palaeophycus heberti?) Bilinichnus simplex Paczesna, p. 36, pi. 5.3 [copy Paczesna, 1985, pi. 4.4] (= Palaeophycus heberti?) double crawling trail Fedonkin, pi. 28.1 Bilinichnus simplex Fedonkin, p. 133134, pi. 26.6) (= Didymaulichnus) Beaconichnus darwinum Acefiolaza and Buatois, p. 96, fig. 117 cf. Beaconichnus darwinum Acefiolaza and Buatois, p. 96-97, fig. TH7 [copy Acefiolaza, 1978, fig. 1-11.2] Beaconichnus darwinum Acefiolaza and Buatois, p. 186,fig.6B

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v.?1993

cf. Beaconichnus darwinum Acenolaza and Buatois, p. 186, fig. 5E [copy Acenolaza, 1978, fig. 1-11.2]

.1994

Diplopodichnus biformis Johnson et ah, p, 399-403, fig. 1 fig. 5

71994

Beaconichnus darwinium? Greb and Chesnut, fig. 12b Diplopedichnus Braddy, p. 101 (lapsus calami).

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1995b .1996

Diplopodichnus biformis Keighley and Pickerill, figs. 1H and 2E-F (partim)

Diagnosis. As for the ichnogenus. Specimens. Fifteen slabs containing forty two traces (TB-135-93, TB-146-93, TB-2-95, TB-595, TB-9-95, TB-10-95, TB-11-95, TB-3-96, TB4-96, TB-5-96, TB-7-96, TB-8-96, TB-12-96, TB20-96, and TB-27-96) housed in the Paleontological Collection of the Emporia State University. Description. Closely spaced imprints that form either two parallel epichnial grooves (Fig. 3B), hypichnial ridges (Fig. 2C), or mark the margins of a furrow preserved as a positive hyporelief (Figs. 2A-B and 3C). In the latter case, two moderately wide grooves are preserved on both sides of the trace. Imprints are elongate, perpendicular to the trace axis, tend to form simple and disorganized rows, and range in length from 1.5 to 1.9 mm. Internal width of the trackway is 0.9-4.0 mm and external width is 1.66.8 mm. Traces can be followed laterally up to 61.4 mm. When preserved in positive relief, furrows are about 0.8-1.0 mm deep. Trace course is straight to winding. In one case, a furrow tapers laterally (Fig. 2B). Overlap between specimens produces false branching (Fig. 2B). Additionally, secondary successive branching has been detected (Figs. 2A-B, Fig. 3A). Quality of preservation of the imprints is highly variable. The poorest preserved imprints are associated with segments that exhibit secondary successive branching. These segments also are characterized by being deeper than the other segments of the trace (Figs. 2A-B).

297

Remarks. Diplopodichnus has been recorded exclusively from the Paleozoic, ranging in age from the Ordovician to the Permian (Johnson et ai, 1994; Brady, 1947). However, if Bilinichnus proves to be its junior synonym, the stratigraphie range of Diplopodichnus extends to the Vendian (Palij et al, 1979). With the exception of the Vendian representatives and a tentative recording from Ordovician offshore shelf deposits (Fillion and Pickerill, 1984; Pickerill et al, 1984), all Diplopodichnus occurrences are from nonmarine to marginal marine settings. The type specimens come from Permian eolian deposits of the Coconino Sandstone in Arizona (Brady, 1947). Johnson et al (1994) recorded D. biformis from Ordovician sediments deposited in transitional subaerial to freshwater environments of the English Lake District. Another possible example was reported by Pollard et al. (1982) from Devonian lake margin deposits of the Hornelen Basin, in Norway. Pollard and Walker (1984), and Walker (1985) described it from Devonian lake margin fades from the Midland Valley of Scotland. Acenolaza and Buatois (1991,1993) documented this ichnotaxon from Carboniferous open-lake sediments and a possible example from Permian playa lake fades of Argentina. These authors also noted that paired trails commonly described as Isopodichnus isp. or Isopodichnus osbornei from late Paleozoic gondwanic gladolacustrine deposits (e.g. Glaessner, 1957; Rocha Campos, 1967; Savage, 1971; Guerra-Sommer et al, 1984) should be placed in Beaconichnus darwinum, and therefore now in Diplopodichnus biformis. The environmental setting of the Devonian occurrences recorded in Antarctica by Gevers et al, 1971 is controversial. While Gevers et al (1971), Gevers and Twomey (1982), Bradshaw (1981), and Bradshaw et al. (1990) favored deposition in marine environments, Woolfe (1990, 1993) suggested nonmarine settings. Examples in marginal-marine settings were reported from the Carboniferous of Kentucky by Greb and Chesnut (1994), and from the Permian of New Mexico by Braddy (1995b). However, specimens from Ken-

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LUISA.BUATOIScraZ.

tucky occur in sediments deposited towards the freshwater zone of an estuary, specifically in estuarine channel successions, whereas those from New Mexico were recorded from possibly freshwater, tidal flat fades. Notably, specimens documented in this contribution also come from transitional fluvial/estuarine fades (cf. Larder et al, 1993; Buatois et al, 1997). Therefore, Diplopodichnus seems to range from fully terrestrial to subaqueous freshwater, and transitional nonmarine/marine environments. Specimens described as Bilinichnus simplex by Fedonkin (1985, pi. 26.6) are actually bilobate furrows rather than paired furrows, and should be placed in Didymaulichnus as indicated by Keighley and Pickerill (1996). The occurrence of Bilinichnus simplex recorded by (Paczesna 1985, 1986) from the Early Cambrian of Poland is based on a single spedmen. Examination of the illustration (Paczesna, 1985, pi. 4.4; 1986, pi. 5.3) suggests that the trace is actually a partially eroded, open burrow with thick, lined, more résistent walls, rather than a pair of ridges. Palaeophycus heberti is the most likely candidate, but restudy of the material should be necessary for a more accurate determination.

conichnus. With respect to D. biformis, they concluded that detailed identification of its producer was not possible, although they tentatively suggested trilobites. However, as it was previously stated, recent research suggests that the Antarctic specimens probably occur in nonmarine environments. (Woolfe, 1990, 1993). Accordingly, Diplopodichnus may be regarded as produced by myriapods in fully terrestrial environments or myriapod-like animals in aquatic settings. Here again is another example of the pitfalls of naming traces after supposed tracemakers, espedally with regard to arthropod trackways. Ichnogenus Diplichnites Dawson 1873 Type ichnospedes. Diplichnites aenigma Dawson, 1873, by original monotypy Diagnosis. Simple trackways consisting of two parallel rows of similar tracks, blunt to elongate, dosely and regularly spaced and roughly normal to the trace axis (after Fillion and Pickerill, 1990).

Diplichnites gouldi (Gevers, in Gevers et al., Diplopodichnus is interpreted as a locomotion 1971) Fig. 1A; Figs. 2A-B, D; Figs. 3A; Figs. 5A-B trace (Repichnia). Gevers et al (1971) noted that .1971 Arthropodichnus gouldi Gevers et al, some trails end in pits or rounded terminations, p. 93, p. 19, figs. 3 and 4 (partim) a feature unreported in all subsequent records of .1973 Beaconichnus gouldi Gevers, p. 1002 the ichnogenus with the exception of spedmens .1975 Beaconichnus gouldi Hänzschel, p. mentioned by Pollard et al. (1982) and Braddy W45, fig. 27.1b (partim) [copy of (1995b). These rounded terminations probably Gevers et al, 1971, pi. 19, fig. 3] represent the place where the tracemaker began .1975 Beaconichnus giganteum Gevers in burrowing under the surface (Braddy, 1995b). HMntzschel, p. W45, fig. 27.1d Brady (1947) discussed potential tracemakers for .1981 Diplichnites gouldi Bradshaw, p. 638this ichnotaxon and concluded that Diplopodich639, figs. 5, 6,29-32 nus was comparable to traces formed by modern .1982 Beaconichnus gouldi Gevers and millipedes, particularly diplopods. Johnson et al Twomey, p. 641, fig. 79.3 (1994) also reviewed potential producers, con.1982 Beaconichnus giganteum Gevers and duding that in their case the animal responsible may have been an aquatic myriapod-like form. Twomey, p. 641-642, figs. 79.4 [copy Gevers et al (1971) discussed possible tracemakHäntzschel, 1975, fig. 27.1d] and 79.5 ers of the ichnospedes formerly placed in Bea.1990 Diplichnites Woolfe, p. 305, fig. 6

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.1991

partially washed-out trackways Hocking, fig. 30 .1994 Diplichnites gouldi Johnson et al., p. 399-^103, figs. 1,4 and 5 .1994 Diplichnites gouldi Trewin, fig. 6H non 1995 Diplichnites gouldi form A Trewin and McNamara, p. 194, figs. 18a-b and 19a (= Umfolozia) .1995 Diplichnites gouldi form B Trewin and McNamara, p. 194, figs. 18c-d, 19b-c, and 20 .1995 Diplichnites gouldi form C Trewin and McNamara, p. 194-195, figs. 18e and 19d non 1995 Diplichnites Trewin and McNamara, fig. 21 (= Palmichnium) .1995b Diplichnites Braddy, p. 101, fig. 1.1 non 1995 Diplichnites Wright et al, p. 307, figs. 5, 6 and 8 Emended diagnosis. Symmetrical trackways that consist of two rows of closely spaced, tapered external, elongate, ellipsoidal to circular imprints, oblique to perpendicular to the trace axis. Large internal and external width relative to track row width. Continuous or discontinuous marks are absent. Imprints are not connected and do not form grooves or ridges (modified from Gevers et al, 1971).

299

Specimens. Eighteen slabs containing forty four traces (TB-112-93, TB-118-93, TB-121-93, TB-122-93, TB-135-93, TB-141-93, TB-1-95, TB5-95, TB-8-95, TB-9-95, TB-10-95, TB-71-95, TB-3-96, TB-9-96, TB-18-96, TB-19-96, TB-2096, and TB-23-96) housed in the Paleontological Collection of the Emporia State University. Description. Symmetrical trackways consisting of two rows of small, closely spaced imprints, which display high preservational variability. Most commonly, the trace consists of slightly imprinted, simple and disorganized track rows. Imprints are tapered externally and elongate or, more rarely, ellipsoidal and oriented perpendicular to the trace axis. Imprints range from 0.8 to 2.1 mm long and are spaced by 0.70.9 mm. Trackway course is straight to curved and can be followed up to 116.9 mm. External width ranges from 5.0 to 8.0 mm. Internal width is 2.1-4.8 mm. Preserved as positive hyporeliefs or negative epireliefs. Remarks. Diplichnites gouldi has previously been reported from Ordovician to Permian rocks (Johnson et al, 1994; Braddy, 1995b). The ichnogenus itself, however, ranges from Cambrian to Triassic (Singh and Rai, 1983; Bromley and Asgaard, 1979). Diplichnites gouldi has been documented from transitional subaerial to freshwater environments of the English Lake District

FIGURE 5 (A) Specimens of D. gouldi showing disorganized rows of unconnected imprints. Negative epireliefs. TB-146-93. (B) Detailed view of/), gouldi. Note elongate, unconnected imprints. Positive hyporelief. TB-191-93. Scale bars = 1 cm

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DUlSA.BUATOIS.tfa/.

(Johnson et al., 1994) and from Silurian fluvial fades of Australia (Trewin and McNamara, 1995). As discussed previously, the environmental setting of the specimens described from the Devonian of Antarctica (Gevers et al, 1971; Gevers and Twomey, 1982) is still a matter of debate, with recent work favoring a nonmarine origin (Woolfe, 1990, 1993). Specimens from the Permian of New Mexico were found in tidal flat deposits transitional between marine limestones and nonmarine redbeds (Braddy, 1995b). Examples documented in this contribution occur in Carboniferous sediments deposited at the estuarine/fluvial transition. Diplichnites gouldi seems to range from transitional terrestrial/subaqueous freshwater to transitional nonmarine/marine environments. A detailed discussion of the ichnotaxonomy of Diplichnites and the status of its different ichnospedes is beyond the scope of this paper. The detailed morphology of D. aenigma, the type ichnospedes, is unclear from the original illustration (Dawson, 1973; fig. 3) and the holotype has not been located (Briggs et al., 1979). However, according to the original description, D. aenigma is considerably larger that D. gouldi. D. gouldi also is smaller than D. cuithensis Briggs, Rolfe and Brannan 1979. Furthermore, the latter is characterized by imprint series of at least twenty three (Briggs et al, 1979; see also Briggs et al, 1984). D. gouldi closely resembles D. triassicus (Linck, 1943), but the latter has imprint series of seven to nine (Pollard, 1985). D. minimus Walter 1988 has elongate to scratch type imprints oriented oblique to the trace axis, and seems to display a tendency to form asymmetric trackways. A detailed and comprehensive review of the ichnogenus Diplichnites, including re-evaluation of its different ichnosperies, is still required. Diplichnites gouldi is considered to represent locomotion traces (Repichnia) of arthropods. It originally was attributed to trilobites by Gevers et al (1971), most likely homalonotids or lichnids (Gevers and Twomey, 1982). Subsequent authors (e.g. Bradshaw, 1981), however, compared this

ichnotaxon with D. cuithensis and suggested arthropleurids (myriapods) as potential tracemakers. Johnson et al (1994) also suggested aquatic myriapod-like forms as possible producers. It should be noted that the Triassic ichnospedes D. triassicus is assodated with small spedmens of Cruziana and Rusophycus, and is interpreted as having been produced by notostracan crustaceans (Bromley and Asgaard, 1979; Pollard, 1985). Myriapods were the most likely tracemakers of the Kansas specimens. Acknowledgements LAB and MGM would like to thank the Kansas Geological Survey for field work and laboratory fadlities. This research was done while LAB and MGM were redpients of external grants awarded by the Argentinian Research Council (CONICET). Partial funding for WPL was provided by NSF grant ERA -9405123 and continuing funding from the Research and Creativity Committee of ESU. WPL also acknowledges the significant contributions of undergraduate students at Emporia State University in the collection of samples utilized in this study and to Rich Leiszler for his research project to catalogue the samples. We also would like to thank Simon Braddy and John Pollard for their careful review of the manuscript, Ron Pickerill for providing valuable comments and a preprint of his Ichnos paper coauthored with Dave Keighley, Mark Schoneweis for the drawings, and John Charlton for the photographic work. References Aceñolaza, F. G. (1978). Trazas fósiles de la Formadón Patquía en el Bordo Atravesado, Sierra de Famatina, La Rioja. Acta Geológien Lilloana, 15:19-29. Acenolaza, , F. G. and Buatois, L. A. (1991). Trazas fósiles del Paleozoico superior continental argentino. Ameghiniana, 28:89-108. Aceñolaza, F. G. and Buatois, L. A. (1993). Nonmarine perigondwanic trace fossils from the Late Paleozoic of Argentina. Ichnos, 2:183-203. Archer, A. W., Larder, W. P., and Feldman, H. R. (1994). Stratigraphy and depositional history within incisedpaleo-valley fills and related fades, Douglas Group (Missourian/Virgilian; Upper Carboniferous) of Kansas, U.S.A. In Boyd, R., Zaitlin, B. A., and Dalrymple, R.

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(eds.), Incised valley systems: origin and sedimentary Chiplonkar, G. W. and Badve, R. M. (1970). Trace fossils from sequences. Society of Economic Paleontologists and the Bagh Beds. Palaeontological Society of India Journal, 14: Mineralogists Special Publication, 51:175-190. 1-10. Bandel, K. (1967). Isopod and limulid marks and trails in Dawson, J. W. (1873). Impressions and footprints of aquatic Tonganoxie Sandstone (Upper Pennsylvanian) of Kananimals and imitative markings on Carboniferous rocks. sas. The University of Kansas, Paleontological ContribuAmerican Journal of Science and Arts, 5:16-24. tions, 19:1-10. Demathieu, G., Gand, G. and Toutin-Morin, N. (1992). La Braddy, S. J. (1995a). The Ichnotaxonomy of the Invertebrate palichnofaune des bassins Permiens Provincaux. GeoTrackways of the Coconino Sandstone (Lower Permian), bios, 25:19^5. Northern Arizona. In Lucas, S. G., and Heckert, A. B. Fedonkin, M. A. (1977). Precambrian-Cambrian ichno(eds.), Early Permian footprints and fades. New Mexico coenoses of the east European platform. In Crimes, T. P., Museum of Natural History and Science Bulletin, 6:219and Harper, J. C. (eds.), Trace Fossils 2. Geological Jour224. nal Spedal Issue, 9:183-194. Braddy, S. J. (1995b). A New Arthropod Trackway and AssoFedonkin, M. A. (1985). Paleoichnology of Vendian Metazoa. ciated Invertebrate Ichnofauna from the Lower Permian In Sokolov, B. S., and Iwanowski, A. B. (eds.), The VenHueco Formation of the Robledo Mountains, Southern dian System, Vol. 1. Historic geology and PaleontologiNew Mexico. In Lucas, S. G., and Heckert, A. B. (eds.), cal basis. Nauka, Moscow, pp. 112-116. (English edition, Early Permian footprints and fades. New Mexico 1990, Springer Verlag, Berlin, pp. 132-137). Museum of Natural History and Science Bulletin, 6:101Feldman, H. R., Gibling, M. R., Archer, A. W., Wightman, 105. W. G. and Larder, W. P. (1995). 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